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The Philippine 












A. S. Arguelles, D.Sc, Editor 
Eduardo Quisumbing, Ph.D., Associate Editor 



Manuel L. Roxas, Ph.D.; F. T. Adriano, Ph.D. 

Joaquin Maranon, D.Sc; R. H. Aguilar, Ch.E. 

Patrocinio Valenzuela, Ph.D.; Marcos M. Aucante, Ph.D. 

A. J. Hermano, D.Sc; Felix V. Espino, Ch.E. 

V. Elicano, B.S.; Antonio D. Alvir^ Ph.D.; Jose Feliciano, Ph.D. 

Experimental Medicine 

Daniel de la Paz, M.D.; Arturo Garcia, M.D.; Onofre Garcu, M.D. 

Cristobal Manalang, M.D.; Isabelo Concepcion, M.D. 

H. W. Wade, M.D.; Walfrido de Leon, M.D. 

Clinical Medicine 

Antonio Sison, M.D.; Liborio Gomez, M.D., Ph.D.; H. Lara, M.D. 
Jose Rodriguez, M.D.; Carmelo Reyes, M.D. 


Elmer D. Merrill, D.Sc; E. B. Copeland, Ph.D.; A. F. Fischer, C.E., M.F. 

T. G. Fajardo, Ph.D.; Rafael B. Espino, Ph.D. 

Nicanor G. Teodoro, Ph.D.; Feliciano M. Clara, Ph.D. 

Nemesio B. Mendiola, Ph.D. 



Heraclio R. Montalban, M.A.; Gonzalo Merino, Ph.D. 

Canuto G. Manuel, D.Sc; Manuel D. Sumulong, M.S., D.V.M. 

Lope M. Yutuc, D.V.M. ; Faustino Q. Otanes, M.S. 

Leopoldo S. Clemente, Ph. D. 

H. O. Beyer, M.A.; Ricardo E. Galang, M.A. 



5c ^..^-- 


No. 1, March, 1949 

tissued January 18, 1960] 

CoPELAND, Edwin B. A. D. E. Elmer: Leaflets of Philippine Botany 1 

CoPELAND, Edwin B. Pteridaceae of New Guinea 5 

Six Plates 

Baisas, p. E,, and Adela U. Pagayon. Notes on Philippine mos- 
quitoes. XV. The chaetotaxy of the pupee and larvie of Trip- 
teroides 48 

Four plates 

Paras, Ernesto M. Blood-chemistry studies in leprosy. III. Total 
calciunfi, diffusible and non-diffusible, calcium, albumin and 

globulin 73 

Hamm, Wm. S., and ARTHtJR Avery. Philippine solar salt industry ..- 85 
EsTAifPADOR, EuLOGio P. Studies on Scylla (Crustacea: Portunldie). 

I. Revision of the genus 95 

Three plates 

Manaix), Gtx>RiA D., and Augustus P. West. Analysis and composi- 
tion of Manila elemi Ill 

Book Reviews 121 

No. 2, lune, 1949 

[Itdued July 22, 19&d] 


Gapuz, Rufino B., and Domingo Santiago. A supplementary guide 
for rapid identification of the larvae of Anopheles minimus 
flavirostris (Ludlow) and Anopheles mangyanus (Banks) 127 

Four plates 

Cabrera, Deogracias J. Observations on the mode of action of 

Naganol (Bayer 205) as a Trypanocidal agent 135 

Cruz, Aurelio O. Composition of Philippine singkamas oil from 

the seeds of Pachyrrhizus erosus (Linn.) Urb 145 

Herre, Alb&rt W. C. T. a new labrid and other interesting 

Philippine fish records 149 

YuTUC, L. M., and Herman Sher. Observations on the distribution 
of Trypanosoma evansi in the bodies of rats and guinea pigs 
during different stages of infection 155 

Vakover^scgh, Moric^. Adverbs and prepositions in Iloko 167 

CoPELAND, Edwin Bingham. Aspleniaceae and Blechnace« of New 

Guinea 207 

Slat plates 

Book Reviews 231 

No. 3, September, 1949 

risstted July W, 19501 


Valenzubla, Patrocinio. Jose Kabigting Santos 275 

UicHANCO, Leofoldo B. A revision of the genus Ectenus Dallas, 

with description of a new species (Hemiptera, Pentatomid®) 285 


iv Contents 

Chakraverti, Dhirendra Nath. Anomalous secondary growth in 

some roots of Hydrocotyle asiatica Linnieus 291 

Two plates 

BOEKENOOGEN, H. A. The carotene content of the fruit of Momordica 

cochinchinensis Spreng 299 

EsTAMPADOR, EuLOGlo P. Scylla (Crustacea: Portunidae) II. Com- 
parative studies on spermatogenesis and oogenesis 301 

Fourteen Plates 

Ramiro, Mariano P. Suitability of Manila hemp waste for paper 

making material : 35$ 

Book Reviews 364 

No. 4, Deeemberj 1949 

tissued March, 1951} 


Pesigan, T. P. Marcos A. Tubangui: in memoriam 367 

CONCBPCION, ISABBLO and RosALiNA L. Dee. Human milk studies: 
I. The thiamine content of mature normal milk and beriberi 

milk , ., 373 

YuTUC, L. M, Observations on the prevalence of tabanid Aies 379 

One plate 

CoPEiiAND, Edwin B. Aspidiacese of New Guinea , 389 

Forty-fottr Plates 


The Philippine 
Journal of Science 

Vol. 78 MARCH, 1949 No. 1 


Adolf Daniel Edward Elmer, son of Jacob Van Dyne and 
Alvina Elmer, was born at Vandyne, Wisconsin, June 14, 1870. 
He married Miss Emma Osterman, of Arlington, Nebraska, 
who survives him, in 1902. 

He was graduated from the Washington State College in 
1899, and finished his work for the degree of Master of Arts 
at Stanford University in 1903. Later in the same year, he 
went to the Philippines, as collector of plants for the Bureau 
of Government Laboratories. This connection lasted only until 
1905, when he began his long career as a private collector 
and student of Philippine plants. 

The life of the private collector is at best a hard one. Its 
only probable personal reward is in serving the collector's 
personal interest. With training, skill, judgment, unremitting 
industry, and with luck, he makes a living. Elmer had the 
other qualifications, but at first no luck except his wife. He 
returned from his first field trip, and began the description 
of his collection and the preparation for distribution; and 
a flood, such as Americans had not yet learned to expect 
in Manila, inundated the city, and practically destroyed the 
entire collection. 

Twice in later years, he had to come in for medical treat- 
ment, to return half-cured to finish his field work. Such 
experience weakens a man permanently, and after his Borneo 
trips he was never able to undertake any hard grind of field 

On the other hand, — ^and this will perhaps give an idea of 
what thorough collection in the tropics involves, — on his Apo 
trip he was seven months without seeing a white man, and 
came out in good condition. But the Apo region is unique. 
It is paradise. In those days, the Bagobos could and would 

6961 1 

2 The Philippine Journal of Science ^^*^ 

explore the forest canopy, and do it with skill. A devoted 
botanist did not so much miss the amenities of white society 
when he could talk botany, and with the Bagobos he could do 
this with mutual satisfaction. As Elmer said, "Those Ba- 
gobos are good botanists." And if the force of this statement 
is not easy to grasp, let me testify that I have never known a 
botanist who could distinguish more local plants than could one 
old Bagobo, Angat. Elmer gave him Latin immortality, 
with Solanum Angatii. Naturally, this Apo collection was 
the most successful ever made by anybody in one season in 
the Philippines. 

Elmer was primarily a botanist. He was a collector because 
collection served his kind of botany. Most collectors arrange 
with a staff of specialists to determine and describe their 
plants. Elmer did this with some specialists, but mostly his 
collections were material for his own study. To serve this 
purpose, he made field notes, the most uniformly good and 
ample I have known any collector to prepare. These notes 
were too ample for inclusion on herbarium labels, but were 
full of information which ought to be published. 

To provide for the prompt publication of descriptions of 
his novelties, in one place and under his own control, Elmer 
decided to undertake the publication himself. Thus began his 
Leaflets of Philippine Botany. Among many such personal 
publications, this has been the outstanding success. At first, 
the printing was done by a local printer. As the collecting 
business was still paying an invisible profit for living expenses, 
he cheapened his publication by buying a little second-hand 
press, and became printer as well as collector, author and 

At this point, his luck unsuspectedly reversed. Having a 
press, he did a job of job printing for a friend, who could not 
pay for it. To cancel the bad account, Elmer took in payment 
some valueless mining stock, and chucked it into a drawer to 
be forgotten. In later years, this stock paid in annual div- 
idends many times its original cost; gave the Elmers a part 
of the measure of comfort in which they spent their later 
years; and gave their son, "our little Anton'' when Ilex An- 
tonii was named, — a Stanford education. Years later, another 
plant was named Antonii, this time for Dr. Anton D. Elmer. 

The Leaflets ran their course, ten volumes, but with con- 
tinuous pagination. In them, Elmer published descriptions 
of 1512 new species, besides a few new genera and varieties. 

78,1 A. D. E. Elmer: Leaflets of Philippine Botany 3 

The number of articles was 137, mostly by Elmer, and mostly 
devoted wholly to the description of novelties. In a few 
papers, he treated families or genera, including all known 
Phihppine representatives. 

A number of articles were by specialists, reporting on El- 
mer's collections. Among the authors of such papers are Radl- 
kofer, on Sapindaceae ; Foxworthy, on dipterocarps ; Ridley, on 
Zinziberaceae ; Beccari, on palms; Martelh, on pandans; A. de 
Candolle, on Elaeocarpus ; C. de Candolle, on Piperaceae; 
Kukenthal, on Carex; Ames, on orchids; Prain and Burkill, 
on yams ; Christensen, on ferns ; Stephani, on Hepaticae ; Broth- 
erus, on mosses; Hieronymus, on Selaginella; Zahlbruckner, 
on lichens; and H. and P. Sydow, Rehm, Murrill and Patouil- 
lard, on fungi. 

Several papers are introduced by descriptions of the country 
where he collected, of the people, their life and industry, as 
well as, of course, the vegetation. These are entertaining 
and instructive reading, like Jagor's records of an earlier day, 
and ought not to be lost by the wider public because they are 
sifted into a very technical publication. Literary sparks occur 
elsewhere too; as where, in the desperate search for vaHd 
specific names in large genera with which we are all familiar. 
Elmer named a Ficus for his own ancestor, as F. Adamsii 
(corrected later to F. Adami, "named after the man who first 
recognized the utility of this genus." 

Elmer worked under his own rules. His descriptions are 
all in English, — ^technically, not valid publication. As to type 
specimens, "holotypes,'' which the botanic world has come to 
respect during the years of Elmer's activity, he used to main- 
tain that all specimens of one of his collection numbers were 
co-ordinate, that one specimen of a new species was as much 
a type as any other specimen. At the end, he fell into line, 
with the statement, p. 3831: "The type specimens of my new 
Philippine species of plants discovered and pubHshed in the 
Leaflets of Philippine Botany are preserved in my private type 

This type collection was kept, for safety, in the Philippine 
National Herbarium, in a "fire-proof" wing of the Bureau of 
Science. What may have endured bombardment was burned 
by the Japanese with the help of gasoline. The distributed 
isotypes are the nearest thing to types that now exist. 

Immortality is conferred on collectors by giving their names 
to the novelties they collect. In such biographies as this, it 

4 The Philippine Journal of Science *^^ 

is common to conclude with a list of plants bearing the name 
of the subject. Of course, Elmer curtailed the list of plants 
bearing his name, by almost every paper he himself wrote. 
Of his fifteen hundred new species, at least a tenth would have 
been likely to bear his name if others had described them. 
Almost one-sixth of his novelties described by others do bear 
his name, but this proportion has to decrease with increase 
of numbers, because there can be only one Ficus Elmeri, one 
Eugenia Elmeri, one Quercus Elmeri, In spite of his describ- 
ing the most of his new species, more than 120 have been 
given his name, a number so great that their listing ceases 
to magnify his honor. Genera are Elmeria, changed to 
Adelmeria; Elmerina; Elmerinula; and Elmeriobryum. 

Elmer died of natural causes, in Manila, April 17, 1942, 
soon after the Japanese occupation, in time to escape the evil 
era of the concentration camps. 

In spite of discouragement and difficulty, he had lived as 
he would. We may quote from him a fitting epitaph: 


Edwin B. Copeland 


By Edwin Bingham Copeland 
Of the University of California, Berkeley 


Genus DICKSONIA LUeritier 

Key to the New Guinea species of Dicksonia 

Major rachises black. • ' ---v- ^< ^^^^^^^ 

Primary pinnules up to 16 by 4 cm 1. D, grandis 

Primary pinnules up to 10 by 2 cm la. D. Ledermanni 

Major rachises dark brown. 

Hairs on rachises few or deciduous 2. D, Schlechteri 

Hairs comparatively persistent. 

Ultimate sterile teeth broadly triangular. 

Stout dark bristles mixed with fine reddish 

ones 3. D, Hieronymi 

Hairs uniform 3a. D. sciurus 

Sterile teeth narrow and sharp 4. D, Archboldii 


D, grandis Ros., Fedde's Repert. 5 (1908) 34. 
Mount Gelu, alt. 1,000 m, Werner 79, isotype in Herb. Univ. 
Calif.; Brass 11897, see comment under D. Ledermanni. En- 


D. Ledermanni Brause, Engler*s Jahrb. 56 (1920) 46. 

Ledermann 8H0, Hunstein Peak, alt. 1,050 m, type, not seen. 
Carr 14769 is received as this species, and I accept the identi- 
fication, because Mr. Alston visited Berlin while working on 
this collection. It is a tree fern, instead of an epiphyte as 
stated on the field label of the type; the pinnules are up to 
10 cm instead of 7 cm long; and the rachises are black rath- 
er than dark-brown. Mr. Alston is in general likely to 
see resemblances where I am more impressed by differences, 
but I still accept this identification. However, I cannot dis- 

* This paper includes the report on the ferns collected by the Third 
Archbold Expedition to New Guinea. Previous reports were published 
in the Philippine Journal of Science 7S (1940) 345-357; 457--469; 7S 
(1941) 347-361; 76 (1941) 23-25. 


6 The Philippine Journal of Science ^^^' 

tinguish Carr H769 from the type collection of D. grandis by 
any probably specific difference. Brass 11897 has the black 
axes of D. grandis; its frond, including the stipe, is 105 cm 
long, even smaller than D. Ledermanni. Except for having 
blacker axes and being more naked, it should be the latter 
species; except for stature, it is D. grandis, I believe it to be 
a depauperate form of D. grandis, and must then mistrust 
the distinctness of D. LedermannL As Brause mistrusted the 
field label of Ledermann 8440, so did I assume a misplaced 
label of Brass 11897, — "Rare low epiphyte in mossy forest;" 
but it is most extraordinary if this is a mistake made twice by 
good collectors or with well handled collections. 


D. Schlechteri Brause, Engler's Jahrb. J^9 (1912) 11. 
Schlechter 1715€, Kani Mountains, alt. 1,000 m, isotype in 
Herb. Univ. Calif; Carr 151S3, Papua, alt. 8,500 feet. Keysser 
II 8 (1912), Mount Bolan, alt. 2,400 to 3,000 m, is var. glob- 
rescens Ros., Fedde's Report. 12 (1913) 165, isotype in Herb. 
Univ. Calif.; it may be a distinct species. Endemic. 


D. Hieronymi Brause, Engler's Jahrb 56 (1920) 48; C. Chr., Brit- 
tonia 2 (1937) 282. 

Ledermann 12851, Sepik region, alt. 1,400 to 1,500 m, not 
seen; Brass 4550, Murray Pass, Papua, alt. 2,840 m, det. C. 
Chr.; Brass 9485, 10711, Lake Habbema, alt. 3,225 and 2,800 m. 


P. sciuTus C. Chr., Brittonia 2 (1937) 283. 
Brass 4991, Mount Tafa, alt. 2,400 m. I find on the isotype 
in Herb. N. Y. Bot. Garden a few stouter, longer and darker 
bristles among the otherwise smaller and brighter reddish 
ones. Endemic. 


D. Archboldii Copel., Univ. Calif. Publ. Bot. 18 (1942) 217. 
Trunco teste Brassio 4 m alto sursum 13 mm crasso deor- 
sum crassiore; foliis circa 10, 3.5 m longis, stipitibus 30-40 
cm longis inclusis, lano basale fulvo-cinnamomeo denso cri- 
nito, stipite sursum rhachique setis fusco-cinnamomeis bre- 
vioribus (5-10 mm) obsitis, maximis sparsis delapsis tuber- 

78» ^ Copeland: Pteridaceae of New Guinea 7 

cula relinquentibus ; pinna mediale 65 cm longa, 22 cm lata, 
subsessile, acuminata, subtripinnata, rhachi densissime seti- 
fera setis apicem versus pinnae parvis pallescentibus ; pin- 
nula primaria recte patente, brevi (1-2 mm)-pedicellata, 
acuminata, deorsum imbricante, basi 3 cm lata; pinnulis se- 
cundariis contiguis, brevipedicellatis, apice aciculati-mucronatis, 
circa 6 mm latis, aut omnino glabris aut costa inferne sparsis- 
sime setulosis; segmentis utroque latere usque ad 6, oblique 
positis, anguste decurrenti-connexis, inferioribus furcatis vel 
bis furcatis lobo acroscopico fertile, aliis acute dentif ormibus ; 
soris 1.5 — 2 mm latis, valvis stramineis integris vel irregularit- 
er crenulatis. 

Dutch New Guinea, 9 kilometers northeast of Lake Habbema, 
alt. 2,700 m, Brass 10970, ^'abundant in forest of lower slopes." 

Characterized most clearly by the acute, dentiform, sterile 
ultimate segments. D. sciurus has some resemblance in this 

Genus CYSTODIUM J. Smith 


C. sorbifolium (J. E. Smith) J. Smith, in Hooker, Genera (1841) PL 

Dieksonia papuana F. v. M, Descr. Pap. PL IV (1876) 76. 

King 230, 350, Brass W8i, H70; Carr 1251^2, 127U, all from 
Papua at minor and moderate altitudes; Schlechter 16519, 
1846S, Kaiser-Wilhelmsland. 

Moluccas; Celebes; Borneo. 

Genus CULCITA Presl 

Culeita villosa C. Chr., Brittonia 2 (1937) 283. 

Brass 11589, Bele River, alt. probably 2,200 m. The type is 
Brass i791, Vanapa Valley, Papua, alt. 1,900 m; isotype in Herb. 
N. Y. Bot. Garden. 

Endemic. Related species from Australia and Polynesia to 
Malaya and the Philippines. 

Genus ORTHIOPTEEIS Copeland 

Ithyeaulon Copel., Univ. Calif. Publ. Bot. 16 (1929) 79. 

The type species of Orthiopteris is O. ferulacea (Moore) Co- 
pel., Bishop Mus. Bull. 59 (1929) 14. That of Ithyeaulon is 

8 The Philippine Journal of Science 


/. moluccanum CopeL, since called /. minus (Hooker) C. Chr. 
The supposed essential difference was that Orthiopteris bore 
hairs, and in Ithycaulon the paleae, on the erect caudex. The 
near affinity, shown by the spores and otherwise, was noted 
in the publication of each genus. It has since appeared that 

0. ferulacea is really paleate. This is most obvious on St. 
John 18311, collected in 1937. The black-brown, thick, rigid 
paleae are rather more than 2 mm long, acute, lanceolate, 
forming a small apical tuft. This tuft is more or less con- 
cealed by the stipe bases of most collections, and the paleae 
are usually narrower. I find them narrower still, when pres- 
ent, on the bases of the stipes, where they suggest the bristles 
of Dipteris. With this correction as to Orthiopteris, Ithycau- 
lon, described a few months later, became a synonym. 

Key to New Guinea species of Orthiopteris, 

Pinnules toothed, with undissected middle area. 

Undissected area over 3 mm wide. 

Indusia narrowly cuneate at base 4. O. minor 

Indusia broadly cuneate at base 5. 0. acuminata 

Undissected area about 2 mm wide 3. O. caudata 

Dissection complete, leaving no middle area. 

Segments and winged axes over 0.5 mm wide 2. O. cicutarioides 

Segments and winged axes under 0.4 mm wide 1. O. trichophylla 

1. O. TRICHOPHYLLA Copel. plate 2. 

O. trichophylla Copel., Univ. Calif. Publ. Bot. 18 (1942) 218. 

O. rhizomate suberecto, 1 cm crasso, paleis castaneis glabris 
integris ovatis 1-2.5 mm longis vestito; stipitibus fasciculatis 
usque ad 30 cm altis, fuscis, basi paleatis et squamulatis, sur- 
sum laminaque glabris; lamina usque ad 65 cm longa et 35 cm 
lata, acuminata, basi rotundata, herbacea, quinquepinnatifide 
dissecta, rhachillis tertiae ordinis et ultimis angustissime ala- 
tis, segmentis 1.5-3 mm longis, 0.2-0.3 mm latis, fertilibus 
ad gerendum soros tantum dilatatis; soris vix 1 mm longis, 
0.5 mm latis, vix basi alatis. 

Brass 12027, 15 kilometers southwest of Bernhard Camp, 
Idenburg River, alt. 1,800 m; "very abundant, tufted, terres- 
trial in rain forest." 

More finely dissected than 0. ferulacea and O. cicutarioides; 
the sori hardly winged; probably the most finely dissected 
fern of its size. 

78.1 Copeland: Pteridaceae of New Guinea 9 

2. O. CICUTARIOIDES (Baker) Copel., comb. nov. 

Davallia cicutarioides Baker, Journal of Bot. 28 (1890) 106. 
Ithycaulon cicutarioides Alston, Journal of Bot. 77 (1939) 289. 
/. tenuisectum C. Chr., Brittonia 2 (1937) 285. 

Brass 12239, alt. 1,800 m; Carr 18257, Boridi, Papua, alt. 
1,500 m; Brass 8919, Papua, alt. 700 m, isotype of /. tenuisec- 
tum in Herb. N. Y. Bot. Garden. I depend upon Alston for the 
interpretation of Baker's species. In describing /. tenuisectum, 
Christensen remarked on the resemblance to O. ferulacea. The 
latter is appreciably more finely divided, and thinner; and the 
sorus is in larger part unwinged, although this feature is not 
usually as conspicuous as it was figured by Hooker, Second 
Century of Ferns (1861) PL 64. Endemic. 

3. O. CAUDATA Copel., comb. nov. 

Saccoloma caudatum Copel., Philip. Jour. Sci. SO (1926) 327. 
Ithycaulon caudatum Copel., Univ. Calif. Publ. Bot. 16 (1929) 80. 

Known only by the type. King 462, Hydrographers Range, 
alt. 900 m. Alston, [Journal of Bot. 77 (1939) 289] would re- 
duce this to his /. cicutarioides. Carr 13257, by which I con- 
strue that species, has the tertiary pinnules more deeply and 
narrowly cut, although several times smaller. 

4. O. MINOR (Hooker) Copel. 

0. minor (Hooker) Copel., Genera (1947) 50. 

Brass 129 U, alt. 1,200 m; 12269, alt. 1,700 m. approaching the 
next species; Bamler, Rosenstock Fil. Novog. Exsicc. n. 116, 
Wareo, alt. 600 m. To Malacca and Fiji. 

5. O. ACUMINATA (Ros.) Copel., comb. nov. 

Dennstaedtia acuminata Ros., Hedwigia 56 (1915) 350. 
Ithycaulon acuminatum CopeL, Univ. Calif. Publ. Bot. 12 (1931) 

Known by the type collection, Bamler 139, Sattelberg, alt. 800 
m, isotype in Herb. Univ. Calif. As just noted, Brass 12269 
is very similar. 

This fern is reduced by Alston, [Journal of Bot. 77 (1939) 
289], to Ithycaulon Novde-Guineae (Ros.) Alston, which is Oe- 
notrichia Novae-Guineae Copel. Alston remarks "I have the 
types of both Rosenstock's species." I have only isotypes, but 
have some reason to regard these as more authentic than the 
types, where there is any difference. Ours are the specimens 
retained by Dr. Rosenstock for his own use when he reputedly 

10 The Philippine Journal of Science ^^^^ 

sold the types. They fit the descriptions perfectly, which can- 
not be the case with Alston's types if the latter are congeneric. 
As to this D. (or /, or O.) Novae-Guineae, Rosenstock states: 
"Eine dem Saccoloma moluccanum (Bl.) Mett. gleichende Art, 
jedoch noch mehr zerteilt, und durch das Indusium wesentlich 
von ihm unterschieden. Dies ist breiter als lang und nach un- 
ten nicht verjtingt." It is certainly no Ithycaulon, no Orthiop* 
teris. O. acuminata has the cuneate indusium base typical of 
the genus. Its distinctness from 0. minor is questionable. 

Genus DENNSTAEDTIA Bernhardi 

Key to New Guinea species of Dennstaedtia 

Pinnae and pinnules not articulate. 
Axes not aculeate. 

Veins setose on both surfaces. 

Herbaceous 1. D. penicillifera 

Subcoriaceous 2. D, concinna 

Veins setose beneath, naked above 3. D, magnifica 

Veins naked on both surfaces. 
Kachises naked or glabrescent. 

Axes light-brown 4. D. cuneata 

Axes dark 5. D, Shawii 

Rachises pubescent 6. D, flaccida 

Major axes aculeate. 

Frond tripinnatiiid 7. D. resinifera 

Frond at least quadripinnatifid. 
Lowest pinnules not dwarfed. 

Herbaceous 8. D, novoguineensis 


Ultimate segments acute 9. D, moluccana 

Ultimate segments rounded 10. H, papuana 

Lowest pinnules dwarfed 11. Z>. scandens 

Pinnae and pinnules articulate 12. D, glabrata 

1. 0. PENICILLIFERA v. A. v. R. 

P. penicillifera v.A.v.R., BuU. Jard. Buit. II No. 28 (1918) 17. 
Hypolepis grandifrons Gepp, in Gibbs, Dutch N. W. New Guinea 
(1917) 195. 

Described from Humboldt Bay, alt. 500 feet. With this 
plant, with limited confidence, I identify King 244, from Pa- 

Mindanao, Copeland 1755. Both of these specimens have 
been confused with D. samoensis (Brack.) Moore, which has 
also been reported from New Guinea, 

T8.1 Copeland: Pteridaceae of New Guinea H 

2. D. CONCINNA Ros. 

D. concinna Eos., Hedwigia 56 (1915) 349. 
Known by the original collection, Bamler 1, Sattelberg, iso- 
type, in Herb. Univ. Calif. Its sori are those of Microlepia 
rather than of Dennstaedtia, and Rosenstock himself changed 
the generic name on our label. However, the affinity is with 
species recognized as Dennstaedtia. 

3. D. MAGNIFICA Copel. Plate 3. 

D. magnifica Copel., Univ. Calif. Publ. Bot. 18 (1942) 218. 

Stipite teste Brassio 3.1 m, lamina 3.9 m longis; rhachi cinna- 
monea setis minutis sparsa; pinna visa (non infima) patente, 
sessile, 110 cm longa, rhachi (nisi in sulca) glabra straminea; 
pinnulis^ circa 18 cm longis, sessilibus, acuminatis, recte pa- 
tentibus, vix contiguis; pinnulis" circa 3 cm longis, basi 1 cm 
latis, deinde ad apicem subacuminatum angustatis; pinnulis"^ 
circa 6 mm longis, supra basin cuneatam 3 mm latis, obtusis, 
oblique profunde incisis, firme papyraceis, costis inferne 
sparse setuliferis aliter glabris, superne atroviridibus inferne 
pallidioribus ; soris in sinubus, deflexis, parvis (0.7 mm latis), 
indusii valva interiore paulo minore. 

Dutch New Guinea, 6 kilometers southwest of Bernhard 
Camp, Idenburg River, alt. 1,050 m. Brass 12982, on bank of 

Distinguished from most of the related species by the acute 
to acuminate secondary pinnules. 

4. D. CUNEATA (J. Smith: Hooker) Moore 

D, cuneata (J. Smith: Hooker) Moore, Index (1857) XCVII. 

Schlecter 16606, Kaiser-Wilhelmsland, alt. 200 m. 
Philippines; Malaya. 

5. D. SHAWn Copel. 

D. Shawii Copel., Philip. Jour. Sci. 30 (1926) 326. 

Known only by the tjrpe. Rev. P. C. Shaw, King 491; moun- 
tains behind Taupota. 

i. D. FLACCIDA (Forster) Bernhard 

D, flaccida (Forster) Bernhard, Schrader's Journal "1800" (1801) 
124, PL 1, /. 3. 

Reported from New Guinea; not known to me. 
Pacific Islands. Although the type of the genus, this spe- 
cies is almost unknown. 

12 The Philippine Journal of Science 


7. D. BESINIFERA (Blttme) Mett. 

D. resinifera (Blume) Mett.: Kuhn, Ann. Lugd. Bat. 4 (1869) 290. 

Listed by report; another mysterious species. 

8. D. NOVOGUINEENSIS (Ros.) Copel., comb. nov. 

Dennstaedtia Smithii var. novoguineensis Ros., Fedde's Repert. 10 
(1912) 323. 

Bamler S 49, Sattelberg, isotype in Herb. Univ. Calif.; Cle- 
mens s. n., Morobe, alt. 4,200 feet Differs from D. Smithii 
most essentially in having spiny axes. The two are probably 
not intimately related. 

0. D. MOLUCCANA (Blrnne) Moore 

D. moluccana (Blume) Moore, Index (1857) XCVII. 

Brass 12267, alt. 1,800 m. Keysser 18, received with this name, 
is a distinct, probably undescribed, species. 
Moluccas; farther range questionable. 


Hypolepis papuana Bailey, Queensland Agric. Journal 23 (1909) 

King UW is probably this species, — was so identified by Bai- 
ley, — ^but has a thing inner (extrorse) indusium, which I can- 
not detect on other sheets, apparently of the same species. 
The Bagobos long ago called my attention to the close affinity 
of Hypolepis and some species of Dennstaedtia. Endemic. 

11. D. SCANDENS (Blume) Moore 

D, scandens (Blume) Moore, Parker's Cat. (1858) ; Index (1861) 

Brass 11519 alt. 2,200 m; Bamler s- n. Rosenstock Fil. Novog. 
Exsicc. n. 20J^, Sattelberg, received as D. moluccana. Schlech- 
ter 18945 probably belongs here, but our specimen is defective. 
Malaya to Tahiti. 

12. D. GLABRATA (Cesati) C. Chr. 

D. glabrata (Cesati) C. Chr., Index (1905) 217; Dansk Bot. Ar- 

kiv. 9 No. 3 (1937) 42. 
Dicksonia glabrata Cesati, Rend. Ac. Napoli 16 (1877) 28. 
Dicksonia rhomhifolia Baker, Journal of Bot. 28 (1890) 105. 
Dicksonia erythrorachis Christ, Ann. Jard. Buit. 15 (1897) 86, 

PL 13, f. 7. 
Dennstaedtia articulata Ros., Fedde's Repert. 10 (1912) 322. 
D. Rosenstockii v.A.v.R., Bull. Jard. Buit. II No. 7 (1912) 11. 

w,i Copeland: Pteridaceae of New Guinea 13 

These synonyms apply to New Guinea specimens, and are ta- 
ken from Christensen, 1. c. (1937). If the species be thus 
broadly construed, I believe that it must include D. ampUi (Bak- 
er) Bedd, also reported from New Guinea. My own belief 
is that several closely related species are here combined; but 
it is certain that more species have been described in this 
group than are justified by the number of collections. 

Brass 13462, alt. 850 m. Many previous collections. 

To Fiji and the Malay Peninsula, as here construed. 


Because the assignment of various species to one or the 
other genus is problematical, it would be better if all Micro- 
lepia species were transferred to Dennstaedtia. The discrim- 
ination of the species is also unsatisfactory. Curiously, nei- 
ther the collection under study, nor the greatest previous one, 
Ledermann's includes any Microlepia. 

Key to New Guinea Species of Microlepia 

Bipinnate, most pinnules subentire 1. M, strigosa 

More compound. 

Rachis black 2. M. melanorhachis 

Rachis stramineous to brown. 

Densely setulose, hairs with bulbous bases 3. M, trichosticha 

Hairs without bulbous bases. 

Pinnules over 10 cm long, lanceolate Denn. concinna 

Pinnules relatively broad, mostly shorter. 

Pinnules 10 by 5 mm, hardly incised 4. M. pseudohirta 

Frond more finely dissected 5. M. Speluncae 

1. M. STRIGOSA (Thunb.) Presl 

M. strigosa (Thunb.) Presl, Epim. (1849) 95. 

A specimen collected by Mrs. Clemens, from Boana, Morobe, 
alt. 1,000 m, seems to be a small form of this species, reported 
from New Guinea, distinguished by small subentire or incised 
pinnules and long, whitish hairs. 

To Japan. 


M. melanorhachis Ros., Fedde's Repert. 12 (1913) 526. 

Keysser 179, Sattelberg Hinterland, alt. 1,400 — 1,500 m, in 
Herb. Univ. Calif. Subquadripinnate, with small, cuneiform 
segments. Endemic. 

14 The Philippine Journal of Science ^^^ 


M, tHchosticha J. Smith, Journal of Bot. S (1841) 416, nomen, 
Davallia trichosticha Hooker, Sp. Fil. I (184-6) 183. 

Cited from New Guinea as var. glabrata Prantl. As a Phil- 
ippine species, this plant is well characterized by dense, short 
hairs with bulbous bases. King 458, Hydrographers Range, 
alt. 600 m, is like it in aspect but not in pubescence. 

Dennstaedtia concinna Eos. bears sori which mostly fall short 
of the margin, and would therefore be looked for in Microlepia. 


M. pseudohirta Ros., Fedde^s Repert. 9 (1911) 425. 

The type is King S08. We do not have Rosenstock's speci- 
men, but this plant seems to be No. 174 of my set. Brass 5295, 
Maf ulu, Papua, alt. 1,250 m, identified by Christensen, is the 
same as King 174, We do have Bamler 29, distributed as Ro- 
senstock Fil. Novog. Exsicc, n. 90 as M. scaberula, but the name 
changed by Rosenstock on our label to M. pseudohirta; it does 
not fit the description, and is very unlike King 174. It is not M. 
scaberula, but may be responsible for the listing of M. scaberula 
as a New Guinea species. Its indusium is drawn upward (to- 
ward the margin) at the sides, and would justify its descrip- 
tion as a new species. 

Schlechter 17575 is also misnamed M. scaberula; it may be 
a large M. pseudohirta. 

5. M. SPELUNCAE (L.) Moore 

M. Speluncae (L.) Moore, Index (1857) XCIII. 

This spcies, regarded as very variable in dissection and pu- 
bescence, is the dumping ground for all Microlepia not named 
otherwise. Several such specimens are in hand from New 
Guinea. Among them may be the M. pilosula listed by Brause. 

All warm lands; in most places a ^'collective species.*' 

Genus OENOTRICHIA Copeland 

O. NOVAE-GUINEAE (Ros.) Copel. 

O. novae- guineae (Ros.) Copel., Univ. Calif. Pub. Bot. 16 (1929) 

Davallia Novae-Guineae Ros., Fedde*s Repert. 5 (1908) 36. 

Known only by the type collection, Warner 78, Mount Gelu, 
alt. 1,700 m, isotjnpe in Herb. Univ. Calif. See comment above, 
under Orthiopteris acuminata. 

'^'* Copeland: Pteridaceae of New Guinea 15 

Genus LINDSAEA Dryander 

Many Lindsaea species which are bipinnate in full develop- 
ment produce also fully fertile simply pinnate fronds. If the 
frond is bipinnate, the leaflets (pinnae) on the main axis, and 
on its branches (technically, pinnules) are alike. In the key 
below I call them all pinnae, and describe the technically bi- 
pinnate fronds as fronds with branched axes. The basiscopic 
margin of the pinna is almost always entire; statements as to 
margin or dissection apply therefore to the acroscopic side. 
This is the only genus of considerable size in which I recognize 
fewer species than Brause did nearly thirty years ago. 

Key to New Guinea species of Lindsaea 
Pinnae dimidiate. 
Veins free unless in the sorus. 
Rachis of frond unbranched. 

Rhizome elongate, fronds distant. 

Stipe short, 5 to 20 mm long. 

Pinnae not deeply incised. 

Larger fronds fully 4 cm wide 1. L. sessilis 

Fronds 2 cm wide 2. L, Foersteri 

Pinnae dissected 3. L. hymenophylloides 

Stipe normally at least 5 cm long. 

Pinnae deeply incised 4. L. roemeriana 

Pinnae crenate or lobed. 

Pinnae cartilaginous-bordered 10. L. marginata 

Pinnae not bordered. 

Pinnae bifid, distal lobe broad ^ 11. L. blanda 

Lobes or crenations several. 

Fronds over 2.5 cm wide 12. L. rhombifoliolata 

Fronds not over 2 cm wide. 

Pinnae about 5 mm wide 8. L. adiantoides 

Pinnae about 3 mm wide 9. L. gracilis 

Rhizome short. 

Pinnae entire to shallowly incised. 
Lowest pinnae crowded, much reduced. 13. L. brevipes 
Lowest pinnae not crowded. 

Stipes brown 14. L. concinna 

Stipes black 15. L. crassipes 

Pinnae dissected. 

Sori widened across segment tips 5. L. Bakeri 

Sori cup-shaped, narrow 5a. L. Schlechteri 

Rhachis branched. 
Rhizome short. 

Pinnae dissected 6. L. tenuifolia 

Pinnae lobed or incised. 

Sorus marginal 16. L. tricrenata 

Sorus inframarginal 17. L. Kingii 

16 The Philippine Journal of Science ^^^ 

Rhizome elongate. 

Pinnae dissected 7. L. Versteegii 

Pinnae entire to shallowly incised. 

Coriaceous 18. L. rigida 


Sori several, submarginal 19. L. microstegia 

Sori few, marginal 16. L. tricrenata 

Veins more or less anastomosing, 
Kachis unbranched. 

Pinnae about 2 mm wide 20. L, sinuato-crenata 

Pinnae over 4 mm. wide. 

Entire plant dark 22. L. obscura 

Plant not remarkably dark. 

Pinnae cartilaginous-bordered 23. L. Schultzei 

Cartilaginous border not evident. 
Rhizome elongate. 

Margin inciso-crenate 24. L. Werneri 

Margin lobed 30. L. stolonifera 

Rhizome short 31. L. decomposita 

Rachis branched. 

Pinnae entire or nearly so. 

Pinnae broadly rounded at apex 25. L. azurea 

Pinnae narrowed to obtuse apex 26. L. papuana 

Pinnae crenate or shallowly incised. 

Basal branches of frond forked 27. L. furcata 

iBasal branches not forked. 

Sori near ends of pinnae 28. L. sepikensis 

Sori along upper and distal margins. 21. L. decomposita 

Pinnae lobed half-way to costa 29. L. davallioides 

Pinnae not dimidiate. 

Frond mostly pinnate, pinnae narrow 31. L. ensifoUa 

Frond mostly bipinnate, pinnae broad 32 L. heterophylla 

1. L. SESSILIS Copel. 

L. sessilis Copel., Philip. Jour. Sci. 6 C (1911) 82. 

Brass 88 W, HoUandia, alt. 50 m; King 2UU (type) and 271 
(juvenile), Brass 1077, 5471, Carr 11992, Papua; Werner 70, 
Keysser, Rosenstock Fit. Novog. Exsicc. n. 215, Kaiser-Wilhelms- 
land. Some of these have been determined also as L. repens, 
L. macraeana, L. pectinata and L. Merrillii, any of which ex- 
cept the last may be a preferable name, according to the fine- 
ness with which one elects to discriminate between species. L. 
sessilis is the common local species or form of the group or spe- 
cies ranging from Mauritius to Hawaii, of which the oldest 
specific name is L. repens (Bory) Thwaites. 

Solomon Islands. 

^^'^ Copeland: Pteridaceae of New Guinea 17 


L. Foersteri Ros., Fedde's Repert. 12 (1913) 527. 

Keysser 193, Sattelberg Hinterland, isotype in Herb. Univ. 
Calif. Like L. sessilis but much smaller; it is approached by 
Brass 5471. 


L. hymenophylloides Blume, Enum. (1828) 218. 

Listed by Brause ; I have no New Guinea specimen. 
Malaya; Philippines. 


L. rocmeriana Ros., Nova Guinea 8 (1912) 719. 

Von Roemer s. n., 731, 1137, 1088, Dutch New Guinea; Brause 
cites three collections by Ledermann. Not seen. We have con- 
strued L. hymneophylloides rather loosely in the Philippines, 
and some specimens could be referred to L. roemeriana. L. 
Wollastonii v.A.v.R. (Odontosoria tenera Ridley) must be some- 
thing of this kind, but its description does not make sense. 

5. L. BAKERI C. Chr. 

L. Bakeri C. Chr., Suppl. Ill (1934) 121; Brittonia 2 (1937) 287. 

Asplenium Bakeri C. Chr., Index (1905) 102. 

Davallia lanceolata Baker, Kew Bull. (1899) 119. 

Lindsaea trichophylla Copel., Philip. Jour. Sci. 6C (1911) 83. 

L. Ledermanni Brause, Engler's Jahrb. 56 (1920) 130. 

This synonymi is by Christensen (1937). 
Brass 13651, alt. 750 m; Lam 825, 944, alt. 80 to 150 m; King 
262 {type of L. trichophylla), Brass SSI 9, Papua. Endemic. 

5a. L. SCHLECHTERI Brause 

L. Schlechteri Brause, Engler's Jahrb. 49 (1912) 28, /. I /. 

Schlechter 19395, Kaiser-Welhelmsland, alt. 250 m. Not seen ; 
but, by description and figure, it seems to be a small L. Bakeri. 


L. tenuifolia Blume, Enum. (1828) 219. 

Lindsaea bhimeana (Hooker) Kuhn, Ann. Lugd. Bat. 4 (1869) 277. 

Brass 1S7S7 A, lS75i, alt. 50 and 570 m; common. 

Malaya and Philippines to Fiji. 

After trying to distinguish L. blumeana from L. tenuifolia, 
and finally deciding that it is impossible, I awaken to the fact 
that Davallia blumeana Hooker could not be the name of a 

6961 2 

18 The Philippine Journal of Science ^^*® 

distinct species, because it was originally published as a syn- 

7. L. VERSTEEGH (Christ) V.A.V.R. 

L. Versteegii (Christ) v.A.v.R., Malayan Ferns Suppl. (1917) 206. 
Odontosoria Versteegii Christ, Nova Guinea 8 (1909) 157. 
L. Rosenstockii Brause, Engler's Jahrb. 56 (1920) 128. 

L. Versteegii is typified by Versteeg 1222. Under L. Rosen- 
stockii, Brause lists five collections by Ledermann. I have seen 
none of these, and cannot distinguish the two species by the 

8. L. ADIANTOIDES (Blume) Kuhn 

L. adiantoides (Blume) Kuhn, Ann. Lugd. Bat. ^ (1869) 278. 

Brass IS^Oi, alt. 850 m, typical; 11869, alt. 1,800 m, very 
small, possibly L. gracilis; 10761, alt. 2,750 m, with a discern- 
ible marginal strand; King ^79; Brass 5079, 5122, 8699, alt. 
2,400 m, Papua. 

Described from the Moluccas, and of questionable ocurrence 
farther West; Fiji. 

9. L. GRACILIS Blume 

L. gracilis Blume, Enum. (1828) 217. 

Two collections by Ledermann reported. I have not seen 
this species here unless Brass 11869 be so identified; it fits the 
description approximately, but comparison with Java spec- 
imens makes me regard it as a small L. adiantoides. The two 
are nearly related, not identical. 

Malaya; Mindanao. 

10. L. MAKGINATA Brause 

L. marginata Brause, Engler's Jahrb. 56 (1920) 126. 

Brass 12021, 12213, imSl, 14131, alt. 850 to 1,500 m; described 
from Ledermann 12660 and 1138^; also var. falcata, Brause No. 
1286lf; none of these seen. Near L. adiantoides, but usually 


11. L. BLANDA Mett. 

L. hlanda Mett.: Kuhn, Linnaea 86 (1869) 80. 

Brass 11256, Bele River, alt. 2,200 m; Schlechter 18699, Bis- 
marck Mountains, alt. 1,800 m. 
Described from Java. 

78»i Copeland: Pteridaceae of New Guinea 19 


L. rhombifoliolata V.A.V.R., Nova Guinea i4 (1924) 4. 
Known only by the type collection, Lam 1892 partim, Door- 
man Mountain, alt. 1,450 m; not seen. It might be an unusually 
ample L. adiantoidejs. 

13. L. BREVDPES Copel. 

L. brevipes CopeL, Philip. Jour. Sci. 6C (1911) 83. 
Brass 13652, alt. 750 m; King 237 (type), 266, Brass 3817, 

Reported from Amboyna. 

14. L. CONCINNA J. Smith 

L. concinna J. Smith: Hooker, Sp. Fil. I (1846) 205, PL 61 B. 

Schlechter ld900, Bander, Rosenstock Fil. Novog. Exsicc. n. 
109, Sattelberg., alt. 500 to 900 m. 
Philippines; Borneo; Palau. 

15. L. CRASSIPES Ros. 

L. crassipes Ros., Fedde's Repert. 5 (1908) 36. 

Brass 13203, alt. 860 m; Werner 7k, isotype in Herb. Univ. 
Calif. L. erassipes is not very near to L. concinna, but may be 
a form of L. cultrata. Clemens 6020 in Herb. Arnold Arb. is 
a mixture, the major part being intermediate between L. cul- 
trata and L. crassipes. The former is as yet unknown in New 
Guinea, and may not safely be reported on this evidence. 

Endemic if distinct. 

16. L. TRICRENATA Baker 

L. tricrenata Baker, Journal of Bot. 28 (1890) 106. 

Known only by the original collection, by MacGregor, Mount 

17. L. KINGII Copel. 

L. Kingii Copel., Philip. Jour. Sci. 6C (1911) 83. 

Brass 8838, alt. 50 m; 8929, Cyclops Mountains, alt. 575 m; 
King 241 (type), 280, Papua. 

18. L. RIGIDA J. Smith 

L. rigida J. Smith: Hooker, Sp. Fil. I (1846) 217, PL 63 A. 
L. monosora Ros., Nova Guinea 8 (1912) 720, non Copel. 
L. monocarpa Ros., in C. Chr., Suppl. I (1913) 49. 

Brass 11761, 12473. L. rigida was previously reported from 
New Guinea, — four collections by Ledermann. L. monosora 

20 The Philippine Journal of Science ^^® 

was based on v. Roemer 785, 1163, not seen. With the mate- 
rial now in hand the supposed distinctions disappear, except 
that I have never seen anastomosing veins in L. rigida. 
Malacca, Ponape. 

19. L. MICROSTEGIA Copel. 

L. microstegia CopeL, Philip. Jour. Sci. ^C (1911) 83. 

Brass 122H, alt. 1,750 m; King 272, type, Papua; Schlechter 
17133, Kani Mountain Range, alt. 1,000 m, received as L. re- 
pens; Bander 126, Sattelberg, received as L. pectinata var. bre- 
vipinnula Ros; Lam 1327, Doorman River, alt. 300 m, received 
as L. pectinata. The Brass and Bamler collections have the 
margin shallowly lobed, one sorus on each lobe. If any New 
Guinea species may be L. repens or L. pectinata, it is not this 
one, but L. sessilis. 


2«. h. SmUATO-CRENATA v.A.v.R. 

L. sinuatO'Crenata v.A.v.R., Nova Guinea 14^ (1924) 30. 

Known only by the t3T)e, Lam 1888, Doorman Mountain, alt. 
1,450 m; not seen. 

21. L. DECOMPOSITA Willd, 

L* decomposita WiUd., Sp. Plant. V (1810) 425. 

Brass 13204, 13415, 134W; very common. King 3S1, 416 have 
smaller and crowded pinnae ; they may represent the ill known 
L. ambigua Cesati. 

India to the Marquesas. 

22. L. OBSCURA Brause 

L. obscura Brause, Engler's Jahrb. 56 (1920) 132. 

Known only by the original collection, Ledermann 8969, Eta- 
penberg, alt. 850 m; not seen. 

23. L. SCHULTZEI Brause 

L. Schultzei Brause, Engler's Jahrb. 49 (1912) 29. 

Brass 11868, alt. 1,800 m, identification not positive. The 
type is Schultze 304 g, Sepik region; not seen. 


A cartilaginous strand reinforces the basiscopic margin in 
many or most species of Lindsaea. It is not rare on the ra- 
chiscopic margin. Extended around all margins, it character- 

^8»i Copeland: Pteridaceae of New Guinea 21 

izes L. rigida, L, marginata and L. Schultzei; but a faint border 
of this kind can sometimes be detected on other species. 

24. L. WERNERI Ros. 

L. Werneri Ros., Fedde's Repert. 5 (1908) 37. 

Brass 13387, alt. 850 m; Werner 5, Mount Gelu, alt. 1,000 m, 
isotjrpe in Herb. Univ. Calif., Schlechter 181U, Finisterre 
Mountains, alt. 950 m. 

A Sumatra specimen, Winkler 194, may belong here. 

25. L. AZUREA Christ. 

L. azurea Christ, Verh. Nat. Ges. Basel 11 (1896) 429; Ann. Jard. 
Buit. 15 (1897) 101, PI U, /. 12. 

Brass 13654, alt. 750 m, typical except for being larger; Brass 
3989, Papua, alt. 50 m. The New Guinea plants should be var. 
Mambae v.A.v.R., Bull. Jard. Buit. 7 (1912) 21, the type of 
which is King i8 p. p. 


26. L. PAPUANA Copel. 

L. papuana Copel., Philip. Jour. Sci. 7C (1912) 68. 

Known only by the type. King 358, Lakekamu, Papua. As 
King sent this plant to me not long before he sent to Buiten- 
zorg the plant named L. azurea var. Mambae, it may be sus- 
pected that they are the same. The brief varietal description 
of the latter might apply to L. papuana. The specimens cited 
above as L. azurea are not L. papuana, even though they ap- 
proach it in size of pinnae. 


27. L. FURCATA Copel. Plate 4. 

I/, furcata Copel., Univ. Calif. Publ. Bot. 18 (1942) 218. 

Synaphlebium, rhizomate 2-3 mm crasso, brevi-repente, paleis 
minutis lanceolatis vestito ; stipite 25-30 cm alto, brunneo, basi 
paleis paucis angustis 2 mm longis sparso, alibi nudo; lamina 
30-35 cm longa, 25 cm lata, bipinnata, pinnis sessilibus, infimis 
typice angulo angusto furcatis, ramo basiscopico 15 cm, acros- 
copico 25 cm longis, pinnis sequentibus uni-jugis 20 cm longis, 
foliolo terminale fere 30 cm longo, pinnis omnibus valde acu- 
minatis ; pinnulis infimis flabellif ormibus circa 7 mm longis, me- 
dialibus dimidiatis, 13 mm longis, basi cuneata 5 mm latis, 
brevissime stipitulatis, patentibus vel erecto-patentibus, apice 

22 The Philippine Journal of Science ^^^ 

rotundatis, herbaceis, superne obscure viridibus margine acros- 
copico leviter crenatis lobis 4 ad 6 truncatis; venis laxe anas- 
tomosantibus ; soris dilatatis, indusio cum margine contermi- 

Dutch New Giiiea, 4 kilometers southwest of Bernhard Cam]), 
Idenburg River, alt. 900 m. Brass 13229, **frequent low epi- 
phyte in rain forest;" also Tto. 13731, ibidem, alt. 850 m, slightly- 
smaller and with the basal pinnae not forked. 

The frond form of the type is like that of Pteris qtuidriaurita, 
with single long basal prongs on the basal pinnae. I regard 
this as the normal frond of the fully mature plant, but it may 
possibly be only an abnormality. Without this peculiarity, the 
species seems to be distinguished by its few long, long-acum- 
inate, ascending pinnae, many ascending pinnules, sparsely anas- 
tomosing veins, and marginal sori. In describing this species, 
I conform to usage and call the branches of the frond pinnae. 

28. L. SEPIKENSIS Brause 

L. sepikensis Brause, Engler's Jahrb. 56 (1920) 131. 

Known only by the type, Ledermann 9423, Etappenberg, alt. 
850 m; not seen. 


L. davallioides Blume, Enum. (1828) 218. 

Reported, and probably not uncommon, but we have no spec- 
imen from "continentar* New Guinea. Bamhoer R 13 is from 
Rook Island, alt. 150 m. 

Malaya; Philippines; Caroline Islands. 

30. h, STOLONIFERA Mett. 

L. stolonifera Mett.: Kuhn, Linnaea 36 (1869) 81. 

Described from New Ireland, cited from the New Hebrides, 
and reported from New Guinea. Not seen. 

31. L. ENSIFOLIA Swartz 

L. ensifolia Swartz, Schrader's Journal "1800" (1801) 77. 

King S 17, Brass 6027, Papua; Bamler 143, Sattelberg, alt. 
100 to 200 m. 
To Madagascar and Fiji. 

32. h. HETEROrilYULA Dry. 

L. heterophylla Dry., Trans. Linn. Soc. 3 (1797) 4.1, PL 8, f. 1. 

Reported here, but not seen. 

To Madagascar and New Caledonia. 

^^'^ Copeland: Pteridaceae of New Guinea 23 

Genus TAPEINIDIUM (Presl) Christensen 

Key to the species of Tapeinidium 

Simply pinnate 1. t. longipinnulum 

At least bipinnate. 

Indusium mostly wider than long 3. T. obtusatum 

Indusium as long as wide 2. T. moluccanum 

Indusium much longer than wide 4. T, stenocarpum 

1. T. LONGIPINNULUM (Cesati) C. Chr. 

T, longipinnulum (Cesati) C. Chr., Suppl. Ill (1934) 176. 
Davallia longipinnula Cesati, Rend. Ac. Napoli 16 (1877) 29. 
D, intramarginalis Cesati, ibidem, teste Christensen. 
T, marginale Copel., Philip. Jour. Sci. ^C (1911) 82. 

Brass 8837, 13845, near sea-level; King 283 (type of T. mar- 
ginale), 349, 

2 T. MOLUCCANUM (Blume) C. Chr. 

T. moluccanum (Blume) C. Chr., Gardens' Bull. 4 (1929) 399. 

Davallia moluccana Blume, Enum. (1828) 237. 

Davallia amboynensis Hooker, Sp. Fil. I (1846) 178, PL 56 C. 

Brass 12102, 12958, 1S863, alt. 850 to 2,100 m; King 261; 
Schlechter, 18185. 


T. obtusatum v.A.v.R., Nova Guinea i^ (1924) 52. 

Known only by the type, Lam 1857, Doorman Top, alt. 2,500 
m; not seen. By description, this does not seem very distinct 
from T. moluccanum. 


T. stenocarpum v.A.v.R., Nova Guinea 14- (1924) 52. 

Lam lkU2, ridge near Idenburg River, alt. 1,420 m, isotype 
in Herb. Univ. Calif. Christensen, Suppl. Ill (1934) 176, re- 
duces this to a variety of T. moluccanum. It seems to me to 
be sufficiently distinguished by its narrow, acute segments, as 
well as by its narrow indusia. As to its identity with Davallia 
stenoloba Baker, I have no opinion. 

Brause lists also T. Denhami, which is T. tenue (Brack.) Co- 
pel., a Fiji species, and T. pinnatum (Cav.) C. Chr. As the 
latter is now construed more strictly than when he wrote, I do 
not know what his specimens would now be named. 

24 The Philippine Journal of Science *^^ 


Key to New Guinea species of Sphenomeris 

Larger segments and sori at least 4 mm wide 1. S, retusa 

Segments about 2 mm wide 2. Odontosoria decipiens 

Segments up to 1.5 mm wide 3. S. chusana 

1. S. RETUSA (Cav.) Maxon 

5. retusa (Cav.) Maxon, Journal Wash. Acad. Sci. S (1913) 144. 
Brass 13279, alt 850 m; Bamler, Rosenstock Fil. Novog. Ex- 
sice. n. 1U7. Typical. 

Philippines. Eeported elsewhere. 

2. ODONTOSORIA DECIPIENS (Cesati) Christ (?) 

Odontosoria decipiens (Cesati) Christ (?) 
King 170, Brass 1434, Papua; Schlechter 16315. Lam 1352, 
identified as this species, has much more narrowly cuneiform 
pinnules, like an enormously overgrown S. chusana. Christen- 
sen, Dansk Bot. Arkiv. 9 No. 3 (1937) 44, maintains, in agree- 
ment with Baker, that this is "a large and more compound 
form," of S. retusa. As he had seen the type, I may not dis- 
pute him; but the specimens cited here are not in my opinion 
S. retusa. 

3. S. CHUSANA (L.) Copel. 

S. chusana (L.) Copel., Bishop Mus. Bull. 59 (1929) 69. 

Brass 10797, 11470, alt. 2,200 and 2,750 m; King 184, Carr. 
18957, Papua. 

Madagascar; India; Japan across Polynesia. 

Genus ISOLOMA J. Smith 
ISOLOMA FULIGINEUM Copel., comb. nov. 

Schizoloma fuligineum CopeL, Philip. Jour. Sci. 1 Suppl. (1906) 

252, PL 1 A. 
Nephrolepis Schizolomae v.A.v.R., BuH. Jard. Buit. T (1912) 22. 
Schizoloma Schizolomae v.A.v.R., Malayan Ferns Suppl. (1917) 214. 

Brass 8804, HoUandia, alt. 20 m. 

Philippines; Borneo. 

Christensen, Suppl. Ill (1934) 170, 171, reduces S. fuligi- 
neum to S. ovatum, to which I cannot assent; and makes S. 
Scfiizolomae a variety of S. guirinianum, which may be rea- 
sonable if one knows how to distinguish a variety from a spe- 
cies. The essential difference between them as species is that 
/. fuligineum has all or practically all veins free; I myself 
distrust this as a distinction. 

^^'* Copeland: Pteridaceae of New Guinea 25 

Genus TAENITIS WiUdenow 

Key to New Guinea species of Taenitis 

Hardly or not at all dimorphic 1. t. blechnoides 

Conspicuously dimorphic 2. T. requiniana 

1. T. BLECIINOIDE3 (Willd.) Swartz 

r. blechnoides (Willd.) Swartz, Syn. Fil. (1906) 24, 220. 

Brass 13293, 13617, alt. 750 to 900 m; Bamler W 3, Wareo, 
alt. 600 m; King 153, Carr 12665, Papua. 
To Assam. 

2. T. REQUINIANA (Gaud.) Copel. 

T, requiniana (Gaud.) Copel., Univ. Calif. Publ. Bot. 16 (1929) 85. 

Bamler, Rosenstock Fil. Novog. Exsicc. n. 252, Rook Island. 
Admiralty Ids.; Melanesia. 

Genus SYNGRAMMA J. Smith 

Key to New Guinea species of Syngramma 

Frond pinnate 1. s. pinnata 

Frond simple 2. S. Hookeri 

1. S. PINNATA J. Smith 

S. pinnata J. Smith, London Journal of Bot. 4 (1845) 168, PL 7, 5, 
/. C. 

Brass 8999, Hollandia, alt. 100 m; King US, 298, Brass 5571, 
Carr 12663, Papua. 
Queensland; Fiji. 

2. S. HOOKERI C. Chr. 

S. Hookeri C. Chr. Index (1905) 346; (1906) 629. 

Brass 12211, 13364, 13369, 13661, alt. 750 to 1,750 m; King 
355, 480, Brass 3804, 5231, Papua. 
Borneo; Fiji, 


In establishing species in this genus, I have treated size, 
form, number and texture of leaflets, their insertion (stipitate 
or decurrent), the number of rows of areolae, and the color of 
the stipe, — all as specific characteristics; and by these have 
thought to distinguish five species, to which Christensen has ad- 
ded a sixth. This may be correct; or, a considerable part of 
the supposed differences may be matters of variation within 
the species. Our collections are too few to afford a basis for 
confident judgment. Following my own precedent, I would now 

26 The Philippine Journal of Science ^^^^ 

describe one or two more species. However, with only seven 
New Guinea collections in hand; looking like five species, it is 
safer to deal tentatively with them, and let the future show 
which differences fluctuate and which are characteristic. 

Key to New Guinea species of Craspedodictyum 

Lateral leaflets decurrent 1. C grande 

Lateral leaflets stalked 2. C. Schlechteri 

1. C. GRANDE CopeL 

C. grande CopeL, Philip. Jour. Sci. 6C (1911) 84. 

King 117, Ambasi, type of genus and species. 

Solomon Islands. 

This plant may be too near to C. quinatum (Hooker) Copel. 
I suppose that Kajewski 508, from Vanikoro, is topotypic and 
typical of that species. The lateral pinnae are decurrent, as 
described by Hooker; my key, Philip. Journal Sci. S8 (1929) 
147, is incorrect ; no New Guinea specimen matches C. quinatum, 

2. C. SCHLECHTERI (Brause) Copel. 

C. Schlechteri (Brause) CopeL, Philip. Jour. Sci. S8 (1929) 147. 

Syngramma Schechteri Brause, Engler's Jahrb. i9 (1912) 

Schlechter 1816i, Finisterre Mountains, alt. 1,000 m, isotype 
in Herb. Univ. Calif. This plant is approximately matched by 
Brass 8932, Cyclops Mountains, alt. 450 m, and 13616, Idenburg 
River, alt. 750 m ; and by Brass 1494, Owen Stanley Range, Pa- 
pua, alt. 1,200 m. To this species, if to either, must be re- 
ferred Brass 13662, Idenburg River, alt. 750 m, and King 268, — 
these two probably identical; and two Bamler collections, Ros- 
enstock Fit. Novog. Exsicc. n. 114 and 114a, presumably col- 
lected together and representing the local range of variation, 
but distributed with different names. 

Genus LEPTOLEPIA Mettenius 

L. Novae-Zelandiae (Col.) Kuhn, Chaetopterides (1882) 348. 

Zahn (1903) Yabim. With a single specimen, two fronds, 
I do not know whether or not the local plant is characterized 
by minor differences from those of Queensland (of which we 
have a single specimen) and New Zealand. However that is, 
if our data of origin are dependable, this is an interesting proof 
of geographic affinity. 

^^'^ Copeland: Pteridaceae of New Guinea 27 

Genus HYPOLEPIS Bernhardi 

Key to New Guinea species of Hypolepis 

Frond three to four times pinnate. 
Rachis smooth or rough, not muricate. 
Axes reddish. 

Axes hairy beneath 1. H, punctata 

Axes more or less naked beneath 2. H, bamleriana 

Axes stramineous or buff. 

Sori abundant 4. H, tenuifolia 

Sori few and one side of pinnules 4a. H, grandifolia 

Rachis spinescent 5. h, papuana 

Frond bipinnate 8. H, Archboldii 

1. H. PUNCTATA (Thunb.) Mett. 

H, punctata (Thunb.) Mett.: Kuhn, Fil. Afric. (1868) 120. 

Bamler, Rosenstock Fil. Novog. Exsicc. n. 173, quite like For- 
mosa specimens; Rrass 102&8, Win, near Lake Habbema, alt. 
2,800 m, scrambling 3 m high, these representing var. obscura 
Brause, Engler's Jahrb. 56 (1920) 161, as does also Brass i970, 
Mount Tafa, Papua. Keysser II 7, Bolan Mountains, alt. 2,400 
to 3,800 m, received from Rosenstock as H. bamleriana /. mi- 
nor, may be a small form of H, punctata. Brass i889. Mount 
Albert Edward, is similar, but bears remarkably long reddish 
hairs beneath the costae. 

New Zealand to Africa, Japan and Hawaii. 


H. bamleriana Ros., Fedde's Repert. 10 (1912) 325. 
Bamler 89 (1908), Logaueng, alt. 300 m, isotype in Herb. 
Univ. Calif. 

3. H. ARCHBOLDII Copel. Plate 5. 

H. Archboldii Copel., Univ. Calif. Publ. Bot. 18 (1942) 218. 
H. distanti Hooker affinis pilosa, rhizomate late repente, gra- 
cile, pilis aurantiacis vestito; stipitibus remotis, 15-20 cm al- 
tis, gracilibus, purpurascentibus, sparse pilosis; fronde 20-30 
cm longa, 5-10 cm lata, acuminata, basi tripinnatifida, rhachi 
pilis fusco-aurantiacis crassis basibus bulbosis vestita; pinnis 
suboppositis, subsessilibus, inf erioribus remotis brevioribus, me- 
dialibus usque ad 5 cm longis basi 15 mm latis, acutis, rhachibus 
deorsum angustissime sursum late alatis, pilis pallidioribus ob- 
sitis; pinnulis oblongis, obtusis, infimis pinnatifidis sequentibus 
lobatis, crenatis et integris, papyraceis, utraque facie sparse 

28 The Philippine Journal of Science ^^^ 

setosis, decidue ciliatis; soris lobos et dentes fere complentibus, 
vix margine reflexo protectis vel omnino nudis. 

Dutch New Guinea, 7 kilometers northeast of Wilhelmina 
Top, alt. 3,560 m, Brause & Myer-Drees 9852, common in 
ground moss in subalpine forest. 

Similar to H. distans, and thus presenting another close link 
with the flora of New Zealand ; and a member of the widespread 
group of H. punctata. Brass 4235, from Mount Albert Edward, 
alt. 3,690 m, determined by Christensen as a variety of H. punc- 
tata, is possibly H, Archboldii, but bears longer hairs, is more 
harsh, and not evidently ciliate. Brass 4389, referred to under 
H, punctata, is more distinct. Mrs. Clemens sends from Mt. 
Sarawaket, alt. 11,000 to 12,000 feet, a minute Hypolepis, the 
fronds 5 to 6 cm long, hardly bipinnate, and fruiting freely. 
It is probably H. Archboldii, and shows how far reduction can 

4. H. TENUDFOLIA (Forster) Bernhardi 

H, tenuifolia (Forster) Bernhardi, Schrade's Neues Journal 1 2 
(1806) 34. 

King 168, Papua. 

New Zealand to Japan. 


H. grandifolia Gepp, in Gibbs, Dutch N. W. New Guinea (1917) 

Gibbs 6258, Humboldt Bay, alt. 150 m; not seen. "The huge 
fronds, the stipitate pinnules, and the few sori distinguish this 
species from the rest," — ^but not very convincingly. 

5. H. PAFUANA Bailey 

H, papuana Bailey, Queensland Agric. Journal 2S (1909) 158. 

Papua, the type collection, of which I have a fragment; also 
King 465, Carr 1222^, Brass 3956. See comment under Denn- 
staedtia, species No. 10. 



H. aculeata Gepp, Journal of Bot. 61 (1923) Suppl. 59. 

Forbes 555, Sogere, Papua, alt. 600 m. I have not seen this 
plant, and cannot by description distinguish it from H. papuana. 

Genus PAESIA St. Hilaire 

Key to New Guinea species of Paesia 

Sterile segments over 1 mm wide, sori long 1. F. radula 

Sterile segments under 1 mm wide, sori short 2. P, lamiana 

'*'* Copeland: Pteridaceae of New Guinea 29 

1. p. RADULA (Baker) C. Chr. 

P. radula (Baker) C. Chr., Index (1906) 476. 

Brass 4603, 4709, Murray Pass, Papua, alt. 2,840 m, det. C. 
Chr. ; Keysser II 59, Bolan, alt. 2,400 to 3,000 m. The latter 
is the collection responsible for the report of P. scaberula, Fed- 
de's Repert. 12 (1913) 162. 



P. lamiana v.A.v.K., Nova Guinea 1-4 (1924) 35. 

Brass 12342, 12U72, alt. 1,650 and 2,150 m. The type, not 
seen, is Lam 1934, Doorman Mountain, alt. 2,480 m. The Brass 
specimens are smaller than the type, and are accordingly at 
most quadripinnatifid, instead of quadripinnate or quinquepin- 
natifid. They seem well distinguished from P. radula by finer 

Species of Paesia have been described from scanty material 
on the apparent assumption that they vary within narrow 
limits. I suspect that they are both variable and plastic, and 
that Rosenstock's identification of Keysser II 59 is correct. In 
that case, P. scaberula ranges from New Zealand to Sumatra, 
and P. radula is a synonym. P. lamiana looks distinct, locally, 
but some New Zealand specimens are similarly dissected. 

Genus PTERIDIUM Gleditsch 

P. aquilinum (L.) Kuhn, in v. Decken, Reisen 3^ Bot. (1879) 11. 

Brass 11490, Bele River, alt. 2,200 m; Carr 13228, Boridi, Pa- 
pua, alt. 1,300 — 1,400 m; these are substantially alike, and rep- 
resent Pteris lanigera Blume. 

Brass 8962, Hollandia, alt. 20 m, is less wooly than No. 1H90, 
and may be P. revolutum (Blume) Nakai. We may have here 
two closely related species, altitudinally separate; or one spe- 
cies which responds with observable difference to the local en- 
vironment; or the difference may be mere variation, appearing 

Genus HISTIOPTERIS (Agardh) J. Smith 

Key to New Guinea species of Histiopteris 

Stipule-like leaflets present i 1. H. indsa 

Stipule-like leaflets wanting 2. H, estipulata 

30 The Philippine Journal of Science ^^^ 

1. H. INCISA (Thunb.) J. Smith 

H. incisa (Thunb.) J. Smith, Hist. Fil. (1875) 295. 

Brass imSO, alt. 2,800 m; 12^20, alt. 1,700 m; Clemens 10165, 
Mt. Sarawaket, alt. 3,300 m; Carr 14^885, alt. 1,450 m. King 229, 
396, Papua. Three species may be represented by these col- 
lections, but not even Brass' are sufficiently complete to provide 
a sure idea of the entire frond. His note on No. 12820 shows 
a height of 3 m, and No. 10750 looks like part of a still larger 

Pantropic, and to Japan and New Zealand. 

2, H. ESTIPULATA v.A.v.R. 

H, estipulata v.A.v.R., Nova Guinea 14^ (1924) 25. 

Brass 120m, 12438, alt 1,750 and 1,800 m, epiphytic. The 
type is Lam 1923, Doorman Mountain, alt. 2,480 m, not seen. 

Genus LEPIDOCAULON Copeland 


L. caudatum Copel., Univ., Calif. Publ. Bot. 18 (1942) 218; 
Genera (1947) 60. 

Rhizomate 3-5 mm crasso scandente, paleis castaneis lan- 
ceolato-ovatis 2 mm longis, decidue caudatis vestito; stipite 25 
cm alto basi cum rhizomate continuo, paleis similibus basi aut 
cordatis aut sagittatis vestito, alibi nudo, atrocastaneo nitido; 
lamina 45 cm longa, 40 cm lata, deltoidea, basi tripinnata, pin- 
nisque caudatis; pinnis suboppositis, infimis 22 cm longis basi 
8 cm latis, basiscopice auctis acroscopice paulo angustatis; pin- 
nula infima basiscopica 7 cm longa attenuato-caudata, pin- 
nulis" circa 6-paribus 1 cm longis 2 mm latis adnatis obtu- 
sis; pinnulis pinnarum superiorum remotis, anguste lanceolatis, 
acutis; costis inferne conspicuis pallidis, venis paucis inconspi- 
cuis, oblique marginem versus salientibus, deinde fertilibus mar- 
gini parallelis. 

Dutch New Guinea, 9 kilometers northeast of Lake Habbema, 
alt. 2,800 m. Brass 10714; "mossy forest of slopes; climbing on 
lower trunks of trees." Also, No. 12051, a single frond. Lake 
Habbema, alt. 3,225 m, an epiphyte in edge of forest; more am- 
ple than the type, and reflexed margin (false indusium) better 
developed. This reflexed margin is no more than suggested on 
the type. 

The elongate rhizome with solenostele and the elongate sub- 
marginal sori or coenosori suggest affinity to such ferns as Lind- 

^^'^ Copeland: Pteridaceae of Neio Guinea 31 

saea ensifolia. The nearer affinity, shown best by the rhizome, 
is to Histiopteris, in spite of free veins and different pattern of 
frond. The paleae are true scales, not mixed nor intergrading 
with hairs. The stipe contains a single bundle, horse-shoe- 
shaped in section. A part of the long-caudate tips of the pin- 
nae of the type are short-f astigiate at the very tip. 

Genus PTERIS Linnaeus 

Key to New Guinea species of Pteris 

Veins free (except in sori). 
Pinnate with simple pinnae. 

Stipe and rachis stramineous 1. P, vittata 

Stipe and rachis purplish 2. P. moluccana 

Pinnate with lower pinnae forked. 

Large fern with many pinnae 8. P. papuana 

Small or medium-sized, pinnae few. 
Stipe smooth, usually stramineous. 

Lamina herbaceous , 4. P. cretica 

Lamina coriaceous 5. P. pellucida 

Stipe rough, dark 6. P. scabripes 

Sparingly bipinnate. 

Fronds uniform 7. P. deltoidea 

Fronds somewhat dimorphic 8. P. ensiformis 

Pinnae pectinate or compound. 

Pinnae pectinate on lower side only 9. P. semipinnata 

Pinnae pectinate on both sides. 

Lowest pinnae not exceedingly enlarged. 
Huge ferns. 

Segments 1 cm wide above base 10. P. excelsa 

Segments much narrower 11. P. Gardneri 

Ferns of small or moderate size. 

Apices of pinnae long caudate 12. P. ligulata 

Apices only moderately elongate. 

Segments over 4 mm wide , 13. P. blumeana 

Segments under 4 mm wide 14. P. pacifica 

Lowest pinnae greatly enlarged, fronds tripartite. 
Ultimate divisions confluent segments. 

Sori short, far from sinus 16. P. Schlechteri 

Sori long. 
Large ferns. 

Segments serrate at apex 17. P. Keysseri 

Segments entire 18. P. macrophylla 

Small, lamina about 20 cm long.... 15. P. Brassii 

Ultimate divisions free pinnules 19. P. montis-wilhelminx 

Areolae single along costae 20. P. wallichiana 

Areolae in single rows along costules 21. P. tripartita 

Areolae numerous. 

Frond pinnatifid or pinnate 22. P. comans 

Frond more compound 23. P. Warburgii 

32 The Philippine Journal of Science ^^^ 

1. p. VITTATA Linn. 

P. vittata L., Sp. Plant. (1753) 1074. 

Brass 11665, Balim River, alt. 1,600 m; apparently common 
at minor and midle altitudes. 
Warm parts of the Orient, and adventitious elsewhere. 

2. P. MOLUCCANA Blume 

P. moluccana Blume, Enum. (1828) 208. 

Several New Guinea collections at middle altitudes. 
Moluccas; Solomon Islands (?) 

3. P. PAPUANA Cesati 

P, papuana Cesati, Rend. Ac. Napoli 16 (1877) 25, 29. 

Brass IISU, alt. 2,200 m; 10886, alt. 2,800 m; Schlechter 
16591, alt. 1,500 m. No. 11314 is the larger specimen, and has no 
forked pinnae. About half of those of No. 10886 are forked or 
twice forked. 

P. bambusoides Gepp, from the Arfak Mountains, described 
from a fragment, and P. Braiisei Eos., Taenitis Brausei, Ros., 
Nova Guinea 8 (1912) 730, seem neither to be adequately dis- 
tinguished from P. papuana. 

Endemic; but not surely distinct from P. opaca of the Phil- 

4. P. CRETICA Linn. 

P. cretica L., Mantissa (1767) 130. 

A species or synspecies of world-wide distribution, reported 
in New Guinea, whence I have no specimen. The three fol- 
lowing species are members of its group. 

5. P. PELLUCIDA Presl 

P. pellucida Presl, Rel. Haenk. I (1825) 55. 

Brass 11508, Bele River, is this or an undescribed relative, 
still larger and with extravangantly caudate pinnae. 
Philippines ; Malaya. 

8. P. SCABRIPES Wall. 

P. scabripes Wall., List (1828) n. 94, nomen; Agardh, Rec. Sp. Gen. 
Pteridis (1889) 11. 

A relative or form of P. cretica, listed here by report. 
Range doubtful. 

7. P. DELTOroEA Copel. 

P. deltoidea Copel., Philip. Jour. Sci. ^C (1911) 85. 
Known only by the type, King 242; Papua. 

^ ^ Copeland: Pteridaceae of New Guinea 33 


P. ensiformis Burm., Fl. Ind. (1768) 230. 

Common at minor altitudes. 
Tropics of the Orient. 

9. p. SEMIPINNATA Linn. 

P. semipinnata L., Sp. Plant. (1753) 1076. 

Listed here by report. 
To Eastern India. 

10. p. EXCELSA Gaud. 

P. excelsa Gaud., in Freycinet, Voyage Bot. (1827) 388. 

The New Guinea specimens I have seen with this name more 
probably represent P. torricelliana Christ, closely related but 
probably distinct. 

A clear-cut Hawaiian species, apparently sporadic even into 

11. p. GARDNERI (Fee) Hooker 

P. Gardneri (Fee) Hooker, Sp. Fil. II (1858) 206. 

Brass 11495, Bele River; Bamler, Rosenstock FiL Novog. Ex- 
sice, n, 84, alt. 300 m; Carr 14431, 15435, Papua. Large, like P. 
torricelliana, but with dark axes and much smaller and more 
numerous segments. 

Philippines; Malaya to Tahiti. 

12. p. LIGULATA Gaud. 

P. ligulata Gaud., in Freycinet, Voyage Bot. (1827) 385. 
P. beccariana C. Chr., Index (1906) 593. 

Apparently common at minor altitudes, inconstant in form of 

Reported from the Solomon Islands to Borneo. 

13. P. BLUMEANA Agardh 

P. blumeana Agardh, Rec. Sp. Gen. Pteridis (1839) 22. 

14. P. PACIFICA Hieron. 

P. pacifica Hieron., Hedwigia 55 (1914) 355. 

Both of these have been treated by most authors as forms of 
a pantropic species, P. quadriaurita, also reported from New 
Guinea, — Bamler J^5. I have thought I could distinguish these 
two segregates elsewhere, but in New Guinea they seem to me 

6961 3 

34 The Philippine Journal of Science ^^^ 

15. P. BRASSII C. Chr. 

P. Brassii C. Chr., Brittonia 2 (1937) 287. 
Brass 5650, Mount Albert Edward, alt. 2,680 m. A small 
plant with deltoid fronds, related to the preceding species. 

16. P. SCHLECHTERI Brause 

P. Schlechteri Brause, Engler's Jahrb. 49 (1912) 33, /. 2 B. 
Finisterre Mountains, alt. 1,200 m, Schlechter 18190, isotype 
in Herb. Univ. Calif. 
Mindanao; Negros; Luzon (P. quinquepartita Copel.). 

17. P. KEYSSERI Ros. 

P. Keysseri Ros., Fedde's Repert. 12 (1913) 167. 

Bolan Mountains, alt. 3,400 to 3,800 m, Keysser II 10, isotype 
in Herb. Univ. Calif.; Mt. Albert Edward, alt. 3,680 m. Brass 


18. p. MACROPHYLLA Copel. Plate 6. 

P. macrophylla Copel., Univ. Calif. Publ. Bot. 18 (1942) 218. 

Rhizomate inviso; stipite teste Brassio 1.5-1.7 m alto, valido, 
brunneo vel stramineo, paleis aciculari-linearibus brunneis 
sparsis et nisi in sulcis deciduis ornato, sparse spinulifero; la- 
mina tripartita, segmento (ramo) mediale 80 cm longo 30 cm 
lato bipinnatifido, pinnis basalibus oppositis 65 cm longis 
erecto-patentibus, pinnula infima basiscopica maxima et simili- 
ter furcata, rhachibus majoribus flabis spinulif eris ; pinnis me- 
dialibus segmenti medialis usque ad 18 cm longis, 2 cm latis, 
sessilibus, caudatis, superne atroviridibus inferne brunneis, 
firme subcoriaceis, costis superne dentibus lanceolatis segmen- 
ts oppositis praeditis; segmentis proximis, adnato-confluenti- 
bus sinubus acutis, circa 12 mm longis, basi 4 mm latis, acutis, 
rectis vel subfalcatis; soris a basi fere ad apicem elongatis, 
indusio integro. 

Dutch New Guinea, 9 kilometers northeast of Lake Hab- 
bema, alt. 2,700 m. Brass 10298; ''forest undergrowth; occa- 
sional in wet bottoms.'' 

Related to P. Keysseri, but larger, with smaller, closer and 
entire segments, and with spinulose axes. 


P. montis'wilhelminae Alston, Journal of Bot. 78 (1940) 229. 
Brass 9045, Lake Habbema, alt. 3,225 m, "frequent in moist 
open places in forest. Two or three rather fleshy large spread- 

^'^ Copeland: Pteridaceae of New Guinea 35 

ing fronds on erect stipes 80-120 cm long;'' Brass & Myer- 
Drees 10056, north slope of Mt. Wilhelmina, alt. 3,950 m, fre- 
quent among the stones of lime-stone screes. The type is Ver- 
steeg 2532, Mt. Wilhelmina, alt. 4,000 m, not seen. The podlike 
pinnules suggest those of Onychium. 

20. p. WALLICHIANA Agardh 

P. wallichiana Agardh, Rec. Sp. Gen. Pteridis (1839) 69. 
Listed here by report. 
India; Malaya; reported from Samoa. 

21. p. TRIPARTITA Swartz 

P. tripartita Swartz, Schrader's Journal "1800" (1801) 67. 
Zahn, King, Carr; common at minor altitudes. 
Old-World Tropics. 

22. p. COMANS Forster 

P. comans Forster, Prod. (1786) 79. 

Reported but unknown to me in New Guinea. 
Polynesia, very common. 

23. P. WARBURGII Christ 

P. Warburgii Christ, Monsunia I (1900) 70. 

Brass 13459, alt. 850 m; apparently common throughout New 
Guinea. An isolated species, with broad lobes or pinnae and 
freely anastomosing veins, suggesting Acrostichum. 


Genus HEMIPTERIS Rosenstock 

H. Werneri Ros., Fedde's Repert. 5 (1908) 38. 

Werner 2, Gelu Station, alt. 1,000 m, isotype in Herb. Univ. 
Calif. ; Schlechter 10944, Kani Gebirge, alt. 1,000 m. 
Solomon Islands, Brass 3329. 


C. MACROPHYLLA (Blume) Hieron. 

C. macrophylla (Blume) Hieron, Hedwigia 5r (1916) 291. 

Brass 12940, alt. 1,200 m; Clemens s. n., Morobe, alt. 1,400 
m; this specimen is exactly like the Mindanao plant called C. 
fraxinea var. Copelandi by Christ, and transferred to C, ma- 
crophylla with the same varietal name by Hieronjonus. Carr 

36 The Philippine Journal of Science *^*^ 

35743 matches better the typical C. macrophylla of Java. Var. 
undulata Hieron,, Engler's Jahrb. 56 (1920) 160, was described 
from Schraderberg, alt. 2,070 m, Ledermann 11 8 IS. 
Java; Mindanao. 

Genus ACROSTICHUM Linnaeus 

A. aureum L., Sp. Plant. (1753) 1069. 
King 204, common along coast. 
Pantropic in brackish swamps. 


Key to New Guinea species of Cheilanthes 

Herbaceous, stipe not scaly. 

Margin reflexed 1- <^- tenuifolia 

Margin flat 2. C. javensis 

Coriaceous, stipe paleate. 

Frond 2.5-4 cm wide 3. C. papuana 

Frond 5-10 cm wide 4. C. belensis 

1. C. TENUIFOLIA (Burm.) Swartz 

C. tenuifolia (Burn.) Swartz, Syn. Fil. (1806) 129, 332. 

Brass 8960, HoUandia, alt. 20 m; King 290. I have also two 
collections purporting to be by Zahn, Sattelberg, alt. 900 m; 
one is this species ; the other is more probably C. Sieberi Kunze, 
but I mistrust the data of collection. 

New Zealand to India. 

2. C. JAVENSIS (Willd.) Moore 

C. javensis (Willd.) Moore, Index (1861) 244. 

Notholaena hirsuta (Poiret) Desv., Journal de Bot. Appl. (1813) 

King 289, Papua. 

Malaya ; Philippines ; Polynesia. 

Notholaena distans R. Br., of New Zealand, Australia and 
New Caledonia, has been reported from New Guinea and 
from Celebes. 

3. C. PAPUANA C. Chr. 

C. papuana C. Chr., Dansk Bot. Arkiv 9 No. 3 (1937) 57. 

Brass 9147, Lake Habbema, alt. 3,225 m; Brass & Myer- 
Drees 9767, 9965, 10057, 10339, Mt. Wilhelmina, alt 8,500 to 
3,960 m, det. C. A. Weatherby. The type is from Mt. Cars- 

'^^' ^ Copekind: Pteridaceae of New Guinea 37 

tenz, 1. Boden Kloss, not seen. Doctor Weatherby's comment 
follows : 

"No. 10s 3 9 is rather strikingly different in appearance from the rest, 
with longer, more remote pinnae, but in all technical characters agrees 
with them. 

"The excellent series obtained by Brass and Myer-Drees shows well the 
range of variation in the species and makes possible some additions to 
and emendations of Doctor Christensen's description. The rhizome is, as 
he inferred, ascending, short and about 5 mm in diameter, very densely 
covered with scales like those of the lower part of the stipe, and producing 
rather numerous, closely approximate fronds. The lower pinnae are not 
reduced; and the indusium, which is very narrow, scarcely 0.2 mm wide, 
is not intramarginal, as described. It may appear so when folded under 
the reflexed margin, but is actually a direct extension of the epidermis, 
apparently of the upper surface. The lower surface of the lamina in 
young fronds is densely beset with small, sessile, pale glands. The spores 
are densely short-echinate with conical, acute, rather thick processes. 

"The collectors' notes describe the lower surface of the lamina in Nos, 
914Tf 9965 and 10057 as "glaucous-white;" no such coloration appears in 
the dried specimens." 

4. C. BELENSIS Weatherby 

C. belensis Weatherby, Univ. Calif. Publ. Bot. 18 (1942) 218. PL 7, 

Rhizoma breve circa 3 cm longum diametro 3 mm in spec- 
imine unico viso adscendens, paleis fusco-castaneis subrigidis 
concoloribus nitidis lineari-lanceolatis acuminatis integris e 
cellulis elongatis utrinque truncatis luminis coloratis vix 
translucentibus compositis dense onustum, apicem versus 
frondes plures laxe caespitosas emittens. Frondes (juveniles) 
and 3.6 dm altae, vetustiores verisimiliter majores. Stipes cras- 
sus teres castaneus opacus leviter albo-ceraceus quam lamina 
multo brevior, paleis brunneis ovatis acuminatis subtenuibus in- 
tegris e cellulis brevibus irregularibus compositis plus minusve 
dense tectus. Lamina elliptico-lanceolata pinnato-pinnatifida 
vel bipinnato-pinnatifida ad apicem pinnatifidum acutum et ad 
basem angustata. Rachis stipiti similis paleas minores spar- 
sioresque gerens. Pinnae ad 16-jugae (in frondibus juvenili- 
bus solum perfectis visis) sessiles oblongae vel oblongo-lanceo- 
latae obtusae latere basiscopico modice producto inaequilaterales 
profunde pinnatifidae vel pinnatae, pagina superiore glabrata 
venulis vix impressis, inferiore plus minusve albo-ceracea, cos- 
tis costulisque prominentibus castaneis. Segmenta lineari- 
oblonga obtusa integra vel | ad costam obtuse lobata, margine 
hyalino angustissimo continuo integro plus minusve glanduloso 
anguste revoluto instructa. Venae subremotae plerumque 2- 
furcatae, venulis sub angulo late acuto abeuntibus, apice cla- 

38 TJ^^ Philippine Journal of Science ^^® 

vellato-incrassato juxta marginem soros gerentes. Sporangia 
in soris singulis pauca brevistipitata, annulo subangusto ce- 
Uulis transverse angustis circa 20. Spori sphaeroidei brunnei 
diametro circa 35 fx rugis crassis subtortuosis processibusque 
late conicis acutis valde asperatis. 

Dry, bare face of limestone cliff; one specimen only. 18 
kilometers northeast of Lake Habbema, Bele River, alt. 2,200 
m, November, 1938, Brass 11210, type in Herb. Arnold Arbore- 
tum, deposited at Gray Herbarium. 

Cheilanthes belensis apparently has no near relatives except 
C. papuana C. Chr., with which it agrees in characters of scales 
and spores and its little developed "indusium." It differs in 
its oblong or oblong-lanceolate pinnae, the lower two or three 
pairs more or less reduced, in its ceraceous lower surface, the 
rachis and costae without capitate glands, and with far fewer 
and smaller scales, and in the non-impressed veins of the up- 
per surface of the lamina. C. farinacea, which, at first sight, 
it resembles, has the scales of the rhizome and stipe with dark 
central band and pale margin, the lower pinnae not reduced, 
the hyaline margin strongly developed, and differently sculp- 
tured spores. 

Genus PELLAEA Link 
P. ZIPPELn (Miqnel ) Baker 

P. Zippelii (Miquel) Baker, Synopsis (1874) 477. 

Originally described from "Nova Guinea, in rupibus.'' 
Schlechter, Engler's Jahrb. 49 (1912) 5, lists this fern as a 
characteristic plant of bare slopes on the Finisterre Moun- 

Endemic, but closely related to P. falcata, and thus another 
witness of affinity to the flora of New Zealand. 


Key to Neiv C^nea species of Doryopteris 

Frond trifid to axis or nearly so.. 1. D. concolor 

Frond cut to a broad wing 2. D. papmna 

1. D. CONOCOLOR (Langsd. and Fischer Kiihn 

D. concolor (Langsd. & Fischer) Kuhn, in v. Decken, Reisen 3 3 Bot. 
(1879) 19. 

Reported and doubtless here, but I have no specimen. 
All warm lands. 

^'»^ Copeland: Pteridaceae of New Guinea 39 

2. D. PAPUANA Copel. 

D. papuana Copel., Philip. Jour. Sci. ^C (1911) 86. 
King 208, type, Goodenough Bay; 388. 

Endemic. I cannot agree with Tryon in reducing this to 
D. ludens. 

Genus ONYCHIUM Kaulfuss 

O. siliculosum Desv. C. Chr., Index (1905) 20; (1906) 469. 
King 209, 333, Papua. 
Malaya; Philippines. 

Genus ADIANTUM Linnaeus 

Key to New Guinea species of Adiantum 
Frond simply pinnate. 
Glabrous 1. ^. philippense 

^^i^y 2. A, caudatum 

Frond more compound. 

H^i^y - 3. A hispidulum 

Glabrous (unless upper side of rachis). 

Stipe at most 10 cm long 4. A, Kingii 

Stipe at least 15 cm long. 

Sori orbicular 5. a, neo-guineense 

Sori wider than long. 

Rachis hairy above 6. A. Christii 

Rachis naked 7. a. hollandiae 


A. philippense L., Sp. Plant. (1753) 1094. 

, Schlechter 16359, 16980, alt. 200 m; Clemens (host of a 
fungus), Morobe; King 211, Papua. 
Africa to Polynesia; immigrant in tropical America. 


A, caudatum L., Mantissa (1771) 308. 

Carr 117 A7, Kanosia, Papua, sea-level. 
West to India and Africa. 

3. A. HISPIDULUM Swartz 

A. hispidulum Swartz, Schrader's Journal "1800" (1801) 82. 
Zahn, Yabim. 
To New Zealand, India and Africa. 

4. A. KINGH Copel. 

A. Kingii Copel., Philip. Jour. Sci. 90 (1914) 5. 
Known only by the type, King 420, Tamata, Papua. 

40 The Philippine Journal of Science 


A, neo-guineense Moore, Gard. Chron. N. S. 7 (1877) 12. 
Known only as from New Guinea, collected by Goldie. 

6. A. CHRISTH Ros. 

A, Christii Ros., Fedde's Report. 12 (1913) 166. 

Brass 10812, 12688. The type is Keysser B 4^, Bolan alt. 
2,400 to 3,000 m, isotype in Herb. Univ. Calif. 


A. hollandiae v.A.v.R., Bull. Jard. Buit. 7 (1912) 1. 

Known only by the type, Gielleriip 9^6, HoUandia, not seen. 
A, fulvum Raoul is also reported, on the cited evidence of 
Ledermann 11761, It is common farther East and South. 


All are photographs of the types 

Plate 1. Dicksonia Archboldii CopeL 

2. Orthiopteris trichophylla Copel. 

3. Dennstaedtia magnifica Copel. 

4. Lindsaea furcata Copel. 

5. Hypolepis Archboldii Copel. 

6. Pteris macrophylla Copel. 


Gopeland: Pteridaceae op New Guinea.] 

[Philip. Jour. Scl, 78, No. 1, 


CopKLAM>: Ptkijidackak of Nia\ (ii'txr.A.l 

IPiiiLir. .loriL St'i.. -78, No. 1. 



fPiULir. .fdriL Sn., 7.H, No. 1. 


Copei.and: FTKK!i»ArKAi% <iF Nkw (JnxiiA.I 

{ .lorn. Sri.. 7H, No. I. 


GOPELANf): I'THlJlIlArKAK «>F Nkw (JriXKA.l iPflllJI'. Jom. SCL, 78, No. 1. 



ToPKI-ANlK I'TKUlDAd'AE OF Nkw (iriXF.A.| 

I l'IlllJl^ JovR. Sc'L, 7S, N«>, 1, 





By F. E. Baisas and Adela U. Pagayon 
Of the Malaria Control Section, Department of Health 


This is an attempt to unify the widely differing terminol- 
ogies used by various authors for the pupal setae of mosquitoes, 
and to harmonize these terms with the popular numeral hair 
notations for the larva. It does seem quite surprising that 
while the pupal hairs are evidently the representatives of 
larval setae, all the schemes heretofore employed for the pupal 
chsetotaxy were conceived in complete disregard of correlation 
or homology with the larval hairs.^ Such a correlation would 
seem naturally indicated when the pupal hairs become suffi- 
ciently visible during development inside the larval skin. In 
available literature and in practice among our own labor- 
atory workers, one of the main criteria to determine a fourth 
instar larva is the appearance of the float hairs. At a certain 
stage these are readily seen through the larval cuticle on ab- 
dominal segment I. Among anophelines, the float hairs are 
directly or nearly directly underneath the palmate hair. These 
are similarly situated in culicines where the palmate hairs are 
represented by what is more commonly known as hair No. 1. 
By a careful examination of suitable specimens, the other pupal 
setae on the abdomen could be seen more or less in the same 
precise location beneath the larval hair to which each corre- 

* Submitted for publication, September 10, 1948. 

^On October 30, 1948, when this was already in press, we received a 
copy of the latest work on pupal chsetotaxy by K. L. Knight and R. W. 
Chamberlain entitled: A new nomenclature for the ch«totaxy of the mos- 
quito pupa, based on a comparative study of the genera. Proc. Helminth. 
Soc. Wash. 15 (1948) 1-10. It was not possible for us to reproduce one 
of their illustrations for comparison with what we show here, or to in- 
clude their scheme in our discussions, but there are fundamental differ- 
ences between their interpretation and numbering of hairs and ours. 
Only No. 5 of abdominal segments IV to VIII, and No. 13 of VIII are 
similarly designated. Their scheme, like others before it, is not based on 
homology or correlation with larval hairs. Of their many illustrations, 
No. 6 represents a pupa of Tripteroides (Mimeteomyia) . 

44 The Philippine Journal of Science ^^^^ 

sponds. The profound changes in the form and contour of 
certain parts when the larva is transformed into pupa nec- 
essarily alter more or less decisively the positions and make- 
up of the hairs on these parts. The correlations of these 
particular hairs are, therefore, extremely difficult to determine. 
But in the main, the ones most valuable from the standpoint 
of classification are not so hard to correlate. 

The most suitable specimens for the study of hair correla- 
tions are those where the pupal setae are sufficiently visible 
through the larval skin. This is usually at the initial stage 
when the larva becomes pale and quiescent, indicating the 
changes that take place during pupation. Though advantageous 
for complete visibility of hairs, a pupa which has started to 
break through the larval skin or when about ready to do so is 
not suitable owing to alterations in position of its hairs. By 
a rapid examination of the living larva under a microscope, 
one can readily determine the suitable specimen. When 
found, it may be killed quickly but not too rapidly in warm 
water. Even in its death throes in warm water, a nearly ma- 
ture pupa may wiggle out of its original position inside the lar- 
val skin. Suitable specimens may be mounted in balsam or any 
other medium containing no acetic acid. Acid clears the im- 
mature pupal hairs to complete invisibility. 

The second half of this paper is likewise an attempt to ho- 
mologize the hairs of a Tripteroides larva with those of an Ano- 
pheles. The schemes promulgated by different workers for the 
hairs on the head of culicines are different from those for ano- 
phelines, and are, moreover, not strictly based on homology. 
Some authors introduced hair notations for the thorax of culi- 
cines similar to the numeral hair designations for the thoracic 
hairs of anophelines, but none has been published to cover the 
abdominal chaetotaxy of culicines. The varying patterns of 
hair arrangement on the head, and the great difference in mor- 
phology from that of anophelines, of abdominal segments VIII 
and IX render the homology of the hairs borne by these parts 
no easy matter to determine. The siphon, particularly, oblit- 
erates all signs of hair correlations. 

In another part of these notes, we intend to include complete 
acknowledgments of what we owe different people concerning 
this study. For the present suffice it to mention those of the 
Department of Health under whom we worked directly, namely. 
Dr. Waif rido de Leon, Chief, Division of Laboratories ; Dr. An- 
tonio Ejercito, Chief, Malaria Control Section; and those of 

^^'^ Baisas and Pagayon: Philippine Mosquitoes 45 

the 19th MGL, U. S. Army, from the Commanding Officer, Col. 
D. M. Kuhns, to the last enlisted and civilian personnel of the 
Entomology Department, who offered to us the use of their 
specimens and laboratory facilities. 

The pupal setse. — Considerable reduction not only in total 
number but also in sizes of individual hairs takes place when 
the larva is transformed into pupa. The relative positions of 
certain hairs are also changed. Fusion of the (larval) head 
and thorax into the so-called (pupal) cephalothorax is accom- 
panied by the elimination of many hairs and the alterations 
in positions of those that are left on these parts. In com- 
parison, the abdominal hairs on I to VII suffer very little 
elimination though their sizes are generally very much re- 

A total of nineteen pairs of hairs arise directly and indirectly 
from the head capsule of a Tripteroides larva; forty two from 
the thorax. But only three pairs of those on the head and nine 
of those on the thorax are represented in the pupal stage. 

In spite of the difference in number of hairs between the lar- 
vsB of Group A and Group B,^ subgenus Tripteroides, the num- 
ber of pupal hairs is exactly the same for both groups. Nor- 
mally each of the species with known pupae of Philippine Trip- 
teroides bears nine pairs of pupal hairs on abdominal segment 
I, eleven on II, while each of III to VII has twelve pairs. 
Quite often the only ventral hair on I and one of the two on 
II are absent; less frequently one of those on VII is also ab- 
sent. Some species seem to have normally only one ventral 
pair even on II. If the socket which Baisas and others^ 
think represents pupal hair 3 (of authors) is included, each of 
segments III to V will have thirteen pairs. Segment VIII 
bears only four pairs of hairs in either group. 

In the larval stage Group B has more hairs on abdominal 
segments I to VII than Group A. This is due to the absence 
of hair 14 in all species of (^roup A. The larva of the local 
Rachisoura does not possess abdominal hairs 2 and 14. A more 
detailed discussion on larval hairs is given below. 

3 Edwards, F. W. Genera Insectorum (932) Fasc. 194. 77-78. 

4 Baisas, F. E. Notes on Philippine mosquitoes, VII. Mon. Bull. Bureau 
of Health, 18 (1938) 182. 

Stone A. and G. H. Penn. A new species of Ctdex from New Guinea 
Jour. Wash. Acad. Sci. $7 (1947) 90, Fig, 7. 

46 The Philippine Journal of Science ^^^ 

A few hairs, in contrast, become more developed in the 
transformation to pupa. The most notable are the superior 
post-ocular hair (No. 3 of Rozeboom & Knight 5), and the 
lateral tufts or the A's of abdominal segments VII and 
VIII. The equivalents of these hairs in the larval stage, as 
we interpret them, are tiny. But the float hair (dendritic 
tuft of authors) which is very well developed and markedly 
exceeds the larval hair it represents in all anophelines and most 
culicines, is rudimentary in all species except one in Group A, 
and though developed in all of Group B is, nevertheless, hardly 
larger than the larval hair to which it corresponds. 

Edwards ^ claims that rudimentary float hair is a general 
characteristic of treehole breeding mosquitoes. But this does 
not seem true for most of the Philippine treehole breeding 
species. Those of the local Megarhinus, Orthopodomyia, Ura^ 
notsenia, and Tripteroides which breed in treeholes have very 
well developed float hairs. Even lesser amount of water in 
narrower spaces than ordinary treeholes does not seem detri- 
mental to the development of the float hairs. The kind of 
Aedes which breeds exclusively in leaf axils of Colocasia; the 
Topomyia found only in very tiny holes of "climbing bamboo" ; 
the Zeugnomyia breeding in the leaf axils of Livistona rotundi- 
folia, the THpteroides known to breed only in pitcher plants; 
and that unusual THpteroides which breeds in very little 
amount of water inside tiny fore3t shells — all of these have 
developed float hairs. 

A few species, however, have rudimentary float hairs. The 
kind of Philippine Ficalbia found in treeholes in jungles, and 
all species, but one, of Group A, subgenus THpteroides, which 
breed in pitcher plants have rudimentary float hairs. 

Differences between the pupsa of Group A and Group B. — 
But for one partially annectant species, the pupse of Group 
A can readily be separated from those of Group B by four 
well marked differences. The annectant form invalidates two 
of these. One is the character of the float hairs, and the other 
is the peculiarity of the respiratory trumpets. The float hairs 
are rudimentary, i. e., single or split into only two to about 
half a dozen branches, and the respiratory trumpets are rela- 
tively more stout with widely expanded but not deeply notched 
apices in Group A (except in the partially annectant species). 

5 Rozeboom, L. E. and K. L. Knight. The Punctulatus complex of Ano- 
pheles. Jour. Parasit. S2 (1946) 128, PL IV. 

6 Edwards, F. W. Mosquitoes of the Ethiopian Region, III (1941). 

78»i Baisas and Pagayon: Philippine Mosquitoes 47 

But in Group B the float hairs are dendritic and the respiratory 
trumpets are less stout and less expanded but deeply notched at 
the apices. The unusual species in Group A has well devel- 
oped float hairs, and very much less stout respiratory trum- 
pets the tips of v^hich are, however, square cut and not 
notched. In common with all species of Group A, this partially 
annectant form has its superior postocular '^ hair comparatively 
much shorter, moderately stout, frayed, single, seldom branched, 
and the paddles do not have any visible midrib. The superior 
postocular hair in Group B is very long and prominent, not 
frayed, single in one species, split into two or three equal 
branches in all the others; while the paddle has very well de- 
fined midrib. 

The hairs of the head. — ^When inside the larval skin, the 
postocular hairs are better seen from the ventral side. These 
are very close to the sites of hairs 13, 14 and 15 on the head 
of the larva. The inferior and median, or what Rozeboom 
and Knight designate as Nos. 1 and 2, are short and slender. 
They are usually not visible because they arise directly beneath 
the anterior portion of the dark imaginal eye. The superior, 
or No. 3 of Rozeboom and Knight, is very much longer spe- 
cially in Group B, and being the only one of the three which 
is truly postocular in position, can readily be seen from the 
dorsal surface. The larval hairs 13, 14 and 15 are weak in 
all known Philippine Tripteroides and usually No. 14 is the 
weakest. By relative positions, the inferior corresponds to No. 
13, the median to No. 15 and the superior to No. 14. No appre- 
ciable change in size of Nos. 13 and 15 takes place in pupation, 
but No. 14 becomes more stout and very much longer. 

The thoracic hairs. — Though very much reduced in number, 
and generally also in sizes, the hairs on the three thoracic 
segments of the larva are represented in the chaetotaxy of the 
pupa. The so-called antero-thoracic pupal hairs,^ or Nos. 4 
to 7 of Rozeboom and Knight, are apparently all that remain 
of the prothoracic larval hairs; the dorsa l and supra-alar, 

7 The term "postocular" is by J. W. S. Macfie in:— The Cha3totaxy of 
the Pupa of Stegomyia fasciata. Bull. Ent. Res. 10 (1920) 161-169. R. 
Crawford uses "ocular" in:— Some Anopheline Pupae of Malaya. (1938) 


8 Knight, K. L. and R. W. Chamberlain consider the antero-thoracic pupal 
hairs part of the mesothoracic setae, but as shown in our Plate 1, fig. 5, 
this is evidently not the case:— Vide— A new nomenclature for the chaae- 
totaxy of the mosquito pupa, based on a comparative study of the genera. 
Proc. Hebnint. Soc. Wash. 15 (1948) 1-10. 

48 The Philippine Journal of Science ^^^ 

Nos. 9 and 10 of Rozeboom and Knight, are the remnants of 
the mesothoracic larval hairs; while the metajiotal (0, P and 
R or 10, 11 and 12 of Rozeboom and Knight) are the repres- 
entatives of the hairs on the metathorax of the larva. 

The arrangement of the four antero-thoracic hairs in their 
original state is shown in Plate 1, fig. 5. The more internal 
two of these hairs are close to each other and farther from 
the other two which are also close together. The innermost is 
either directly underneath or a little anterior or posterior or 
somewhat internal or external to hair 1 on the prothorax of 
the larva. The other is similarly placed in relation to larval 
pro-hair 3. Pro-hair 2 of the larva seems eliminated. 

Though also somewhat variable in position, the most ex- 
ternal of these pupal hairs does not arise (in the Tripteroides 
specimens on hand) outer to the site of larval pro-hair 4. It 
may, however, be a little anterior or antero-internal or pos- 
tero-internal to that hair. If this is taken as the equivalent 
of larval hair 4, the other would have to be considered the 
representative of either the larval pro-0 or 2. No. 2 seems 
out of the question, while is weak and not correlated in 
position to this hair. Our choice is to consider the outer- 
most pupal hair as No. 5 and the other as No. 4, even if there 
is apparently as much reason to consider them No. 7 and No. 5 
respectively since in all species of Tripteroides the larval pro- 
hairs 5 and 7 are developed. But the distance from the outer- 
most pupal hair to the site of larval hair 7 is too wide to be 
in harmony with that which intervenes between the innermost 
pupal hair and the pro-hair 1 of the larva. It would seem 
reasonable to expect the two antero-thoracic pupal hairs to 
be external to larval pro-hair 4 if they were the equivalents 
of the larval pro-hairs 5 and 7. Moreover, the tendency of 
the cephalothoracic hairs to be more internally located than the 
larval hairs they represent is least on the region of the antero- 
thoracic, somewhat more where the dorsal and supra-alar hairs 
are located, and greatest on the metanotal. 

When inside the larval skin or on mounted whole pupae, the 
identity of the four antero-thoracic hairs, according to the no- 
menclature of Crawford or the numbers given by Rozeboom and 
Knight, is not readily evident. Furthermore, the upper of these 
hairs, as seen in quite a good series of mounted pupal skins of 
different Tripteroides on hand, sometimes seem to become the 
lower. Plate 1, fig. 5 shows our interpretation of these and 
other cephalothoracic hairs according to what seem their re- 

^8» 1 Balsas and Pagayon: Philippine Mosquitoes 49 

spective equivalents in the larval hairs. We consider the inner- 
most of the antero-thoracic or the ''upper anterior" of Crawford 
or No. 6 of Rozeboom and Knight, as No. 1 ; the next or the 
"lower anterior" (of C.) or No. 4 (of R. & K.) as No. 3; the 
third or the "upper posterior" (of Crawford) or No. 5 (of 
Rozeboom and Knight) as No. 4; and the fourth or outermost 
or the "lower post»a.rior" (of C.) or No. 7 (of R. & K.) as No. 5. 
The dorsal hair (No. 8) and the supra-alar (No. 9) ap- 
parently represent the mesothoracic larval hairs 1 and 5 re- 
spectively. In Group A, No. 8 arises at the same level (under- 
neath) but a little internal to the mesothoracic hair 1 of the 
larva, and the two are nearly equal in rank in most species. 
In Group B this hair is either directly beneath the base of, 
or a little anterior or posterior to, the large stellate tuft (No. 1) 
on the mesothorax of the larva; No. 9 is less obvious in ho- 
mology, it being seemingly equal, in different species, to either 
2 or 3 or 4 of the mesothoracic larval hairs. But as No. 5 is 
one of the most developed mesothoracic setse, we prefer to 
regard the supra-alar as the remains of this hair. 

The metanotal hairs (O, P and R of authors, or Nos. 10, 11, 
and 12 of Rozeboom and Knight) seem to represent respectively 
1, 2 and 5 of the metathoracic larval hairs. The metanotal 
hairs do not diifer significantly between the two groups of 
Tripteroides, but in the larval stage at least No. 1 is a large 
stellate tuft in Group B; much smaller, often single, in Group 
A. No. 12 is the one directly or nearly directly beneath larval 
hair No. 1 ; Nos. 10 and 11 being internal to it and not imme- 
diately near the site of any larval hair. This appears to be 
due to the contraction of the metanotum (metathorax) when 
transformed into pupa. 

The abdominal hairs, — Because there is a maximum of only 
twelve pairs of hairs (thirteen if the socket on III to V is in- 
cluded) on an abdominal segment of a Tripteroides pupa re- 
gardless of the difference in number of hairs between Group 
A and Group B in the larval stage, it seems reasonable to 
assume that the hairs carried over to the pupal stage are 
identical in all species. Likewise (and logically it seems) the 
hairs not found on the larva cannot presumably be expected to 
appear on the pupa specially because, the tendency in pupation 
seems to be the reduction of hairs. But this does not seem to 
be always the case. Hair 2 which is absent from the 
larva of one species, is still represented on the pupa of even 
this species; and though tiny, is not less so than those in 
met — 4 

50 The Philippine Journal of Science ^^*® 

which this hair is well developed in the larval stage. On the 
other hand, No. 14 which is developed in Group B, but absent 
in Group A, is not represented on any of the known pupae of 
Philippine Tripteroides. 

The remarkable similarities in number and general arrange- 
ment of the pupal setae in different genera and subgenera of 
mosquitoes is contrasted more or less markedly by the differ- 
ences in their respective larval chsetotaxy. This would seem 
to indicate that development and retrogression of hairs are 
first manifested in the larval stage, such that "new'' larval 
hairs may not be expected to appear in the pupal stage, just 
as "old" ones which have already been eliminated from the 
larva may still be represented in the pupal stage. Abdominal 
hair No. 2 may be taken as an "old'' hair which has been elim- 
inated from the larva of one species but still exists on the 
pupa of even this species. Apparently this hair is being dis- 
carded from the larva of Group A since it is very tiny when 
present, and often absent from some segments or from all 
segments of some individuals in any species. In contrast, 
this hair is well developed and stellate in the larva of Group 
B, but also very tiny like that in Group A in the pupal stage. 
For a "new" hair. No. 14 seems this kind. It is well dev- 
eloped and stellate in the larva of Group B, but absent in 
Group A, and not represented in the pupal setse of any species. 

The general likeness in arrangement of pupal hairs dis- 
played by different mosquitoes does not strictly mean homo- 
logy between similarly placed hairs. This is because some 
hairs vary in positions not only in different species, but also 
on different segments of an individual, and sometimes also on 
different individuals of the same species. Similarities between 
two or more hairs, and their close proximity to each other on 
any part or segment increase the difficulty of recognizing these 
hairs. Moreover, when the pupa emerges from the larval skin 
and stretches out to assume its normal form, the relative posi- 
tions of some hairs (indicated when yet inside the larval skin) 
are altered. Abdominal hairs 3, 4, 6, and 8 are specially 
affected by this stretching. Only by comparing the setse of a 
free pupa or pupal skin with those of an immature pupa inside 
the larval skin of the same species can these hairs be properly 

Hair B, or what we designate as No. 5, is one of the most 
variable in position in different species and on different seg- 
ments. Normally, in most species this is posterior and in- 

^^'* Baisas and Pagayon: Philippine Mosquitoes 51 

ternal to Nos, 3 and 4 (of our scheme) on abdominal seg^ 
ments and IV. But it is often external to 3 and posterior to 
4 on V to VII; or Nos. 3, 4 and 5 may be evenly aligned one 
after another. In certain species, No. 5 is normally internal 
to 4. 

The arrangement of hairs on VII among individuals of any 
species in Group A is very variable. No. 5 (or B) of this seg- 
ment is often so close to No. 6 (or A) it may be mistaken as 
(our) No. 8; or it may be rather far away and well internal 
to 4. Sometimes it is weaker than either 3 or 4. In mounted 
flat preparations, Nos. 8 and 12 (of our scheme) on this seg- 
ment may appear either dorsal or ventral ; while No. 8 is some- 
times unusually close to 6. 

Two or three species in Group A have all the abdominal hairs 
weak. Even B and C or Nos. 5 and 1 of these species cannot 
be easily picked out unless correlated with the positions of 
the larval hairs. 

Quite often in routine examination, one runs across larvae 
which are ready to molt to the fourth instar. In this stadium, 
each of the fourth instar larval hairs is beautifully placed 
directly underneath the corresponding hair on the enveloping 
third instar larval skin. There can be no question about hair 
equivalents in this. But the picture is not as clearly carried 
over to the stage when the fourth instar larva is about to shed 
its skin and give rise to an entirely different form, the pupa. 
Every part of the larva undergoes changes in pupation, and 
the number and arrangement of hairs are as much affected as 
the degree of change that takes place in the part where such 
hairs are located. It seems the fourth abdominal segment 
suffers the least change in form, and the hairs it bears are 
the least altered in arrangement. From segment IV the 
changes increase progressively either anteriorly or posteriorly, 
the greatest being at segment I, thorax, and head, and at the 
terminal segments of the abdomen. We use segment IV, there- 
fore, as the basis of interpreting hair relationships, but at the 
same time we give due value to relationships which are better 
indicated elsewhere. Correlations between the lateral hairs, for 
example, are better shown on VI and VII (see Plate 1, figs. 
1 and 2). 

In parts or segments where changes due to pupation are 
great, homology of pupal hairs cannot be strictly based on prox- 
imity to certain larval hairs. For instance, on segment VII 
as shown in Plate 1, fig. 6, the pupal hair that corresponds to 

52 The Philippine Journal of Science ^^*^ 

larval hair 5 is directly underneath larval hair 4, while that 
which represents larval hair 4 is directly under larval hair 
3. The true relationships of these hairs can be correctly un- 
derstood only from hair correlations indicated on the middle 

Among the few Anopheles and Culex pupae we examined, the 
large pupal hairs B and C on the abdomen are readily recog- 
nized since they are, on most segments, directly beneath lar- 
val hairs 5 and 1 respectively. In Tripteroides, however, B 
is more or less ventral with respect to larval hair 5, and C 
directly or almost directly under larval hair 1. 

On the ventral side the relationship between pupal and 
larval hairs is less apparent. If judged only by its position 
on IV, pupal hair 8 (of authors) would seem to correspond 
to larval hair No. 11. But hair 8 is evidently identical with 
the only ventral hair (often absent) on I, which in turn ap- 
pears to represent hair 13 of the larva. This varies in pos- 
ition on different segments and is unrecognizable when all the 
ventral setae are of about the same sizes or types. The other 
ventral hairs, D, 6 and 7 (of authors) seem to be the equiv- 
alents of larval hairs 9, 12 and 7 respectively. D or 9 is 
usually the longest; 7 ranks next and may equal or even ex- 
ceed 9 on the more posterior segments; while 12 is about 
equal to 13. As No. 12 becomes dorsal in position on V-VII 
in Group A, only three pairs are on the ventral surface of 
each of these segments except on VII in cases or individuals 
where what we propose to be designated No. 8 becomes ven- 
tral, or where No. 8 is dorsal, but 12 is on the ventral side. 
Mounted flat preparations are often confusing since some of 
the dorsal and ventral hairs come to equally clear definition 
under the same focus either under high or low magnifications. 

The pair of tiny ventral hairs (No. 9 of authors) found in 
all Anopheles and most culicines is absent in Tripteroides. The 
pair on VIII of Tripteroides being similar to 9 on VIII of 
other culicines may be mistaken for this hair. But what Trip- 
teroides possesses, as such, seems more reasonably the equiv- 
alent of No. 8 (of authors) on the preceding segments, or what 
we consider No. 13. Hair 13 is present also on VIII of the 
larva among Tripteroides, although this is laterally located in 
Group B; ventral, as usual, in Group A. 

The chsetotaxy of segments I and II differs somewhat from 
that of III-VII. H and the float hairs, just like C and 
C of II-VII arise directly underneath or closely underneath 

'S' 1 Baisas and Pagayon: Philippine Mosquitoes 53 

larval hairs and 1 respectively, and so the float hair should 
be designated No. 1, and H, No. 0. K, L, M, S, T, and U are not 
immediately near the location of larval hairs. K in most spe- 
cies in Group B is the most prominent seta outside of the 
float hair, but it is comparatively much weaker in species of 
Group A. It seems the equivalent of what we regard as hair 
2 (or 5 of authors) on the following segments. " This may 
appear questionable, first, because hair 2 is not present on 
segment I of the larvae among Tripteroides, and, second, it 
greatly outranks 2 of the other segments. Perhaps this is a 
vestige of an ancestral form which had hair 2 on I as in 
anophelines, while its size is understandable in the light of 
unusual variations in sizes of hairs. The Ih on I and II in all 
kinds of mosquito larvse are a good deal larger than those 
of other segments, yet in the pupal stage these are very much 
reduced on I and II and well-developed on VII and VIII. L, 
M, Sy T, and U seem to represent larval hairs 3, 4, 5, 7 and 6, 

In the nomenclature of authors for anophelines, segment II 
of the pupa is credited with two pairs of No. 1 and two pairs 
of No. 2 hairs. Even when in its original position inside the 
larval skin, the homology of the two No. 1 pupal hairs with 
the larval setse is not evident because they are situated and 
pressed close to pupal hair A (of authors) on the lateral side 
of the segment. When released from the larval covering, they 
become farther apart. It seems that the first or the more 
external corresponds to the long secondary lateral hair or No. 
7 of the larva, while the other corresponds to another lateral 
larval hair, No. 8. The more posterior, but often the more 
internal of the two No. 2 pupal hairs arises closely beneath 
larval hair No. 4, and the other closely beneath larval hair 
No. 3. B on this segment is rather far from larval hair No. 5, 
so its homology is based on evidences manifested on other 
segments. C and C of segment II, like those of the succeeding 
segments, arise directly under or closely beneath larval hairs 
1 and 0, respectively. 

There is usually one pair of ventral hair on I and two on II. 
.However, that on I and one of the two on II are absent from 
some individuals of species in which these hairs are normally 
present. Some species evidently have normally only one pair 
even on II. The ventral hairs are variable in position accord- 
ing to species and somewhat variable also among individuals 
within a species. That on I appears to be the equivalent of 


The Philippine Journal of Science 


ttarval hair 13. When only one pair is on the ventral side of 
II it seiems to be also No. 13; and when there are two ventral 
pairs, the more external and anterior in position appears to 
be the equivalent of larval hair 12 ; No. 13, the inner and more 

As a summary, and to harmonize the pupal hair notations 
with those of the larva, the following changes are proposed: 
The post-ocular hairs. 

The inferior (of Macfie), or No. 1 (of Rozeboom and Knight), should 
be No. 13 to correspond to sub-basal or post-mandibular hair or 
No. 13 of the larva. 
The median, or No. 2 should be No. 15 to correspond to the infra- 
orbital hair, or No. 15, of the larva. 
The superior, or No. 3, should be No. 14 to correspond to the orbital 
hair, or No. 14, of the larva. 
The antero-thoracic hairs. 

The upper-anterior, or No. 6, should be pro-1 to correspond to pro- 
thoracic larval hair No. 1. 
The lower-anterior (of authors), or No. 4 (of Rozeboom and Knight), 

should be pro-3 to correspond to prothoracic hair 3 of the larva. 
The upper-posterior, or 5, (sometimes this is the lower) should be 

pro-4 to correspond to prothoracic larval hair No. 4. 
The lower-posterior, or 7 (sometimes this is the upper) should be 

pro-5 to correspond to larval hair No. 6 of the prothorax. 
The dorsal hair, or No. 8, should be meso-1 to correspond to hair 

No. 1 on the mesothorax of the larva. 
The supra-alar hair, or No. 9, should be meso-5 to correspond to hair 
No. 5 on the mesothorax of the larva. 
The metanotal hairs. 

O, F, and R, or Nos. 10, 11, and 12, should respectively be meta-1, 
meta-2, and meta-5 to represent the larval hairs on the meta- 
thorax of the same designations. 
The abdominal hairs. 

Proposed designations 

according to homology or 

apparent homology with 

larval hairs of the same 


Designations by authors. 

Designations by Rozeboom 
and Knight. 

H and C 

9 a nd 5 


Float hair and C 


2 _,___. 

iCandS _ 

4 on I, 7 on other segments 


L, anterior 2 of II, and 4 of other 

5 on I, 6 on other segments 


M, posterior 2 of II, and 2 of other 


5 _ > 

S. B and A' 


A and U -.. _. 



T, first No. 1 of II, and 7 of other 


g ^ 

Second 1 of II and 1 of other seg- 


9 . _ > 



10 - 

3 (socket on III-V) _. 


12 _ > - 

6 _ 


13 ^ . _ _ 

8- _ 


^8>i Balsas and Pagayon: Philippine Mosquitoes 55 

To regard pupal hairs A and U as the equivalents of larval 
hair 6 or the Ih (as we propose here) may seem questionable 
because, while Ih of I and II are well developed, those on the 
following segments decrease progressively in length such that 
No. 6 on VII (of most species) and on VIII (of all species) 
are very tiny. By contrast, U and A's of the pupa become in- 
creasingly developed from I to VIII, U on I being tiny, but 
A's of VII and VIII are* nearly always large subplumose 
tufts. Perhaps this may be explained by the fact that changes 
in hairs in the process of pupation do not follow the same di- 
rection, and that in part at least, it seems development or re- 
trogression of hairs depends upon the usefulness of such hairs. 
Hair Ih is evidently most useful to the larva on I and II, just 
as the A's are apparently of greatest utility tothe pupa on VII 
and VIII ; hence, the development of Ih in reverse direction to 
that of U and A. Perhaps the marked increase in size of lar- 
val hair 14 when it becomes the so-called superior post-ocular 
hair of the pupa is similar to the change of Ih on VII and on 

The illustrations in this paper were drawn by Mr. Eliseo 
Enriquez, artist of the Malaria Section. These are all orig- 
inal except the one copied from Rozeboom and Knight (for com- 
parative purposes), and that which is reproduced from Baisas 
(1938) which was also originally dra^vn by Mr. Enriquez. 


The numerical notation for larval hairs of Anopheles employed 
by Puri,9 Christophers 10 and Rozeboom and Knight (loc, cit) is 
here adapted for the hairs of Tripteroides larvse. Correlating 
the hairs of Tripteroides with those of Anopheles is difficult 
only on the head and abdominal segments VIII-X. The setal 
arrangement on the thorax and abdominal segments I to VII 
is closely similar to that of anophelines. Barraud^^ illustrates 
and numbers the thoracic chsetotaxy of Tripteroides, From his 
interpretation, however, we differ in the case of metathoracic 
hairs 3, 4 and 5. As shown in Plate 1, fig. 5 of the present 

*PURI, I. M. LarvaB of anopheline mosquitoes with full description of 
those of Indian species. Ind. Med. Res. Mem. No. 21 (1931), Plates VII 
and IX, 

'''Christophers, S. R. Fauna of British India. Diptera. Family Cu- 
licidae. Tribe Anophelini. ^ (1933) p. 39, Fig, 10; and p. 43, Fig, 11. 

"Barraud, p. J. Fauna of British India. Diptera. Family Culicidse. 
Tribes Megarhinini and Culicini 5 (1934) 36, Fig, 9g, 

56 The Philippine Journal of Science *^*® 

paper, we follow the sequence of numbers according to the 
arrangement of hairs displayed on the pro- and mesothoraces ; 
i. e., outwardly and somewhat upwardly from 1 to 4, and down- 
wardly to 5. 

Contrast in hairs between Groups A and B of subgenus THp- 
teroides is very marked. The majority or at least some of the 
main hairs in Group A are spines or spinelike, or else very long 
and stout but single; whereas the* dominant hairs in group B 
are the large stellate tufts. Stellate tufts, when present in 
Group A, are not predominant, and are of different types from 
those of Group B. But apart from the major differences be- 
tween the two groups, there are secondary dissimilarities be- 
tween species in each group which indicate deviations from the 
main direction of hair development. For instance, in Group 
A the shortness of all the hairs, including the Ih of the abdo- 
men in two species, is about as remarkable as the absence of 
spines from the thoracic pleural hair groups in another two 
species, or the very thick abdominal hair 13 in still two other 
species; or the distinct differences in abdominal hair 1 which 
can be used to divide the group into three smaller units. 

Division between Groups A and B, though recognizable by 
the general types of hairs composing the thoracic pleural 
groups, is not clear cut as regards the pleural spines. Such 
spines are present in all but two species of Group A, and ab- 
sent in all but two species and three subspecies of Group B. 
The spines when present are very much thicker in Group A 
than in Group B. 

Four kinds of deviations from the main direction of hair 
development may be recognized in Group B. Most notable is 
the presence of one or two spines on the prothoracic pleural 
plate. New species m^^ jg unique in having its prothoracic 
hairs 9 and 10 modified into spines, and this is the only known 
species which has some of its hairs on the head, less so on the 
thorax and abdomen, conspicuously flattened. The powelli 
complex is also distinct. While in this the prothoracic hair No. 
9 is long and branched as usual for Group B, No. 10 is trans- 
formed into a slender spine — similar to a branch of a large 
stellate tuft though somewhat longer and thicker. A very wide 
range in number of branches of the stellate tufts is shown by the 

12 The new species and subspecies, of which there are twenty one will be 
named and described in subsequent papers. Here these are referred to 
merely by letters. 

^s»i Baisas and Pagayon: Philippine Mosquitoes 57 

powelli complex. The higher number of branches seems asso- 
ciated with dark larvae, while the low counts occur among pale 
individuals. The third type of deviation is represented in new 
species j. It has its prothoracic hair 1 on its own plate sep- 
arate and some distance away from that which bears 2 and 
3. Its Nos. 5 and 6 are branch«3d into two or more ; the meso- 
and the meta-hair 1 are not stellate but tiny with two or more 
branches, sometimes single ; while its meso-13 differs from those 
of all the others in being long and single. Its abdominal hair 
12 is also unusually long and prominent. The fourth type 
which embraces the majority of Group B (perhaps best rep- 
resented by dyari), does not have any spine on the thoracic 
pleural plates, no hairs are distinctly flattened, and the varia- 
tion in number of branches in the stellate tufts does not seem 
as wide as in the powelli complex. 

The larva of the only known species of Rachisoura in the 
Philippines has its own pattern of hair development; is quite 
easily separable from those of subgenus Tripteroides, and dif- 
fers markedly from known larvae of Australasian Rachisoura. 
Its adult is different from all known Rachisoura in being ornate. 

The head. — Assuming, as commonly accepted, that the ar- 
rangement of hairs in Anopheles is the most primitive and 
therefore the most ''fixed," the changes in position of hairs 
among culicines may be better understood from a comparison 
with anophelines. Thus compared, it becomes evident that 
the differences between the hairs on the head of an anopheline 
and a Tripteroides larva are more in types and arrangement 
rather than in actual number. For while an Anopheles has 
twenty one pairs on the head, a Tripteroides has nineteen. On- 
ly the post-clypeal or No. 4, and that which arises near the apex 
of the maxillary palp or No. 16 are not represented in Tripte- 

On the pre-clypeal plate of an anopheline larva are located 
only two pairs of hairs— the inner pre-clypeal or No. 1, and 
the outer pre-clypeal which is not given any numeral desig- 
nation by authors probably because it is not used in classi- 
fication. There are constantly four pairs on the pre-clypeal 
plate in Group A Tripteroides, but there may be four, three 
and a half, or three pairs in Group B. This is because in 
Group B the innermost pair is variable in position, one or 
both members of the pair being on the fronto-clypeal plate 
in some species. Regardless, however, of the instability in 

58 T^e Philippine Journal of Science ^^^^ 

position of certain hairs, the pre- and fronto-clypeal plates 
together bear a total of eight pairs of hairs in all known 
Philippine Tripteroides ; nine in Anopheles. Hence, actually 
only one pair is missing from Tripteroides, Because of the 
evidence of movement or tendency to move indicated by the 
innermost pair, it seems probable that other hairs have also 
moved, but have become fixed in their new positions. Fron- 
to-clypeal hairs A, B and C of culicines, which authorities 
regard as the equivalents of the frontal hairs in anophelines, 
are good examples of those which have assumed different fixed 
positions. From this, it seems reasonable to consider two of 
the pairs on the pre-clypeus of Tripteroides represent two of 
the clypeal hairs in Anopheles. 

The thick preclypeal spines apparently correspond to No. 1 
hair in Anopheles. The long, slender pair directly inner to 
the preclypeal spines in Group A, but variably located above 
or posterior, though also internal, to the spines in Group B, 
appears to be the equivalent of the inner clypeal hairs or No. 
2. Some distance posterior to the preclypeal spines, very close 
to the dividing line between the pre- and fronto-clypeal plates 
are two tiny pairs. Very often they are not visible because 
of the dark background specially when the feeding brushes 
are infolded, and because they are tiny and closely appressed 
(bent downwardly) to th»a preclypeal plate. The outer of 
these two seems to be the equivalent of the outer preclypeal 
in Anopheles, more so because of its similar position, and in 
two species it is flattened like those in many anophelines. The 
inner pair seems to be the representative of the outer clypeal 
hair (or No. 3) of Anopheles. If considered independently 
of anopheline notations, the innermost pair should be desig- 
nated No. 1; the preclypeal spines. No. 2; the inner of the 
tiny pairs. No. 3; and the outer (outermost of the four pairs), 
No. 4. But if homology or apparent homology is given prim- 
ary consideration as it should be, the preclypeal spines should 
be No. 1, the innermost pair No. 2; No. 3 should be as it is; 
and No. 4 may be left as such, for the sake of convenience, 
since it does not seem to represent the post-clypeal or No. 4 
of anophelines. 

While hair A of the fronto-clypeus is fairly fixed in position 
in all culicines we have examined, B and C are not. Among 
those species which have these three hairs in straight line, 
or where the line is not very badly broken, it is not difficult 

^«'i Baisas and Pagayon: Philippine Mosquitoes 59 

to determine their correlation with the frontal hairs of Ano- 
pheles. But where one pair is very much forward or far behind 
in position, the usual method followed by authors is to regard 
the most anterior pair as B, regardless of whether this is in- 
ternal or external to C. This seems erroneous when the most 
anterior pair is also the most internal, because (in Tripte- 
roides at least) the movement of (clypeal) hairs seems forward 
(or backward) instead of inward or outward, although certain 
hairs (the sutural) manifest both types of movements. The 
great majority among known species of Philippine Tripte- 
roides have the most anterior frontal pair also the most in- 
ternal; in one or two this is directly anterior to what in the 
usual method will be considered C. We propose, therefore, to 
regard the most external pair as A; the most internal (in the 
majority of species) as C, and the remaining pair, wherever 
it may be, as B; or No. 5 for C (most internal). No. 6 for 
the next or B, and No. 7 for the most external or A, No. 
5 arises at the* very (anterior) border of the fronto-clypeal 
plate in some species of Group A; a little behind that point in 
others. It is single in some, branched in others. Group B dif- 
fers in having hair 5 much more posteriorly located. No. 6, is 
more or less at the same level with 7 in Group A, usually more 
posterior in Group B. This is fairly long, slender, single in all 
species except in n. sp. j. Hair 8 or the sutural or e of authors 
is much more anterior and internal in position in Group A than 
in Group B; it is even internal though posterior to 6 in n. sp. 
b. It is shorter than the other fronto-clypeal hairs, and 
branched except in two species. 

Hairs on the epicranial plate. — The trans-sutural, or No. 9, 
is similarly located as in anophelines. This is rather weak 
and usually branched, non-diagnostic. No. 12, or the sub-an- 
tennal, hair is more posteriorly located, branched in all spe- 
cies, but very short in Group A; fairly large tuft in Group B. 
The sub-basal or post-mandibular hair (No. 13) is more ven- 
tral, often at the same level with 12. It is usually single, dis- 
tinctly flattened in new species, m. The orbital (No. 14) is 
laterally placed as in anophelines, weak, non-diagnostic. No. 
15, or the infra-orbital, is farther forward, sometimes a little 
away from the anterior margin of the imaginal eye. These 
two hairs are seldom useful in classification. The post-maxil- 
lary hair (No. 18) is very weak, single or spHt into two or 
more branches in Group A, but fairly long in microcala and 

60 r/ie Philippine Journal of Science ^^^^ 

nepenthicola, these two having 13 and 15 stronger than in 
other species of the group. On most species of Group B 
hair 18 is closer to the mentum in position. This is weak 
or moderately large, but very large in two of the undescribed 
species, m and n, where it constitutes the most developed hair 
on the head. The submental or hair No. 20 in Group A is at 
approximately the same location as that of anophelines but it is 
very much posterior, close to the rim of the epicranial plate in 
Group B. It is branched in all species, weak in Group A, slight- 
ly more developed in Group B. 

Other hairs on the head. — No. 10, or that at the tip of the 
antenna, is moderately long and stout, single. No. 11, or the 
shaft hair, arises not far from the apex of the antenna, fairly 
long or moderately short, single. The hair which springs 
from the basal piece of the maxilla, or No. 17, is w^^ak, single 
or branched in Group A; a fairly stout though small tuft in 
most species of Group B. This hair frequently appears as if 
it arises from the base of the maxillary palp because the suture 
dividing the basal piece from the palp is often not well defined. 
Hair 19 is fairly long, stout, usually single in both groups; 
it is the strongest ventral hair in some species of Group A. 
The hair which arises near the apex of the maxillary palp, 
or No. 16, is not present in Tripteroides. 

The thorax. — There are fifteen pairs of hairs on the pro- 
thorax of Tripteroides larva, which conform closely in ar- 
rangement to those of Anopheles and may similarly be num- 
bered from to 14. Only fourteen pairs are on the mesotho- 
rax and thirteen on the metathorax. No. 13 is very weak or 
absent on the mesothorax of two species in Group A, but No. 
14 is present as usual. 

Between Groups A and B, the contrast in hairs of the tho- 
rax is about equal to that of the abdomen, and certainly more 
marked than that of the head. While the tendency of the main 
thoracic hairs of Group A is to become thick spines, or spine- 
like with a few branches, or else very long and stout, but 
single, those of Group B tend to become large stellate tufts 
with many branches. The branches usually vary from about 
ten to over thirty, of fairly uniform lengths. When large 
stellate hairs are present in Group A, the branches are usually 
markedly different in lengths, more stout, taper distinctly from 
about the middle to the tip, the apex being blunt or jagged. 

^8>i Baisas and Pagayon: Philippine Mosquitoes 61 

coarse. The branches of the stellate tufts in Group B are more 
slender, hardly taper toward the apex except of those like No. 3 
of the prothorax which end in finely drawn points. More often, 
however, as in the majority of the tufts, the branches terminate 
into a single or double or triple fairly coarse non-divergent 
points. Fraying or barbs are usual in all the principal hairs 
or spines of both groups. Nos. 0, 3, 4, 5, 7, 13, and 14 of the 
prothorax, Nos. 1, 8, 13, and 14 of the mesothorax, and Nos. 1 
and 13 of the metathorax are large stellate tufts in Group B, but 
are either spines, spinelike, tiny or very long and prominent in 
Group A. No. 7 of the meso- and metathorax are thick spines 
of varying lengths according to species, but that on the meta- 
thorax is normally split into two or more coarse points in Group 
B ; constantly single in Group A. Two new species of Group B 
have the mesothoracic spine (hair 7) rather weak, often 
branched, its small tubercle, unlike those of all other species, 
not fused with the plate of No. 6. On the other hand, the meso- 
spine of new species d is quite long but not very thick, nearly 
like a thick normal hair. All but one species of Group A have 
No. 10 (of the pleural group) on the pro- and meso-thoraces 
modified into thick short single spines; in two species Nos. 9, 10 
and 12 of the three thoracic segments are reduced to short 
thick spines. But new species d and g (Group A) have No. 10 
long and normal. Except in new species m, Group B has No. 
9 long and branched on the three thoracic segments; No. 10, 
outside of the powelli complex, dvari, and two new species, 
is about as long as 9 but single on all segments. No. 12 is 
similar to 10 though usually longer on the pro- and mesothorax, 
but always more slender and very much shorter on the metatho- 
rax. Prothoracic hairs 9 and 10 of an undescribed species 
m, are spines of moderate lengths and thickness, 10 somewhat 
exceeding 9 in both respects. These are single but compara- 
tively much less stout than the pleural spines in Group A. 
These are among the most marked departures from the general 
direction of hair development in Group B, which are equalled 
by the absence of spines from the thoracic pleural hair groups 
in two species of Group A. 

Prothoracic hairs 1 to 3, otherwise known as the shoulder 
hairs, arise from a single common plate in nearly all species of 
both groups. The same is true with 5 to 7. Exceptions to these 
are distinctive specifically. Such as in new species j where hair 
No. 1 is on its own plate separate and some distance away from 

62 The Philippine Journal of Science ^^^ 

that which bears 2 and 3; and in an undescribed species, e, 
which has its hairs 1 to 7, sometimes to 7, on a common con- 
spicuous plate. Two species of Group A have pro-13 and 14 on 
a single plate. Sometimes hair 4 in an undescribed species, d, 
has the tubercle attached to the plate from which 5 to 7 spring. 

On the mesothorax only the plates of 6 and 7 are fused; but 
in an undescribed species j\ these are on separate small tu- 
bercles. Again this species is unique in having meso-13 non- 
stellate, i. e., long and single. Unlike that on the pro- or the 
metathorax, meso-6 is not internal to 7, but closely anterior 
or external to it. 

Hair 6 also arises from the large tubercle bearing the spine, 
or No. 7, of the metathorax in all species of Group A, and in 
the only Rachisoura known locally. The point of attachment, 
however, in Rachisoura (undescribed species, i) is at the very 
base of the tubercle some distance from the base of the spine 
itself. In species where 6 is not so attached, it is also located 
close to this point. 


Hair 0, — Found only on the prothorax, is a tiny tuft with 
from about six to over twenty branches, and situated close to 
the shoulder hairs in Group A ; a large stellate tuft, though not 
quite as large as the other stellate hairs, situated more or less 
directly behind hair No. 4 in Group B. 

No. i.— Spinelike, variable in length according to species, 
single or split into two on the prothorax of Group A; either 
more or less like this on the mjeso- and meta-thoraces, or that 
on the mesothorax is very long, single, or as in new species d, 
this is a tiny tuft on both the meso- and meta-thoraces. In 
Group B, No. 1 is a large stellate tuft on all thoracic segments, 
the branches stiff, terminating in single, double or triple points, 
except in an undescribed species, /, which has No. 1 of the meso- 
and meta-thoraces slender, weak, split into two to about half 
a doz»dn branches, sometimes single. 

No. 2. — Invariably single, slender on the prothorax in all spe- 
cies of both groups; about the same on the mesothorax, but 
branched on the metathorax (except in one species) of Group 
A; nearly the same on the meso- and meta-thoraces of Group 
B, though occasionally branched on metathorax. 

No. 3. — In Group A single, shorter than 2 on the prothorax; 
longer, single or branched on the meso- and meta-thoraces. 

^«'^ Baisas and Pagayon: Philippine Mosquitoes 63 

This is stellate on the prothorax of Group B, often slightly 
larger than 1, and with the branches ending in fine-drawn sin- 
gle points; usually quite long but single on the mesothorax; 
also single but much shorter on the metathorax. 

No. 4.— In Group A single, usually equal in rank to No. 2 on 
the prothorax, about the same on the mesothorax, a tiny tuft 
on the metathorax. In Group B a stellate tuft, like No. 1, on 
the prothorax, stellate but smaller on the metathorax, single, 
slender on the mesothorax. 

No. 5.— In Group A either spinelike, similar to No. 1 or 
much longer, with fine-drawn tip on the prothorax; invariably 
long, single on the mesothorax; but tiny, single, branched on 
the metathorax. In Group B long, slender, single on the pro- 
thorax, similar but longer on the mesothorax, stellate tuft on 
the metathorax. The only exception is an undescribed species, 
j, which has its pro-hair 5 split into fairly stout branches. 

No. 6.— In both groups slender, fairly long, single, on plate 
with, and internal to, 7 on the prothorax; closely anterior or 
slightly external to, and on plate with, 7 on the mesothorax; in- 
ternal to and on plate with 7 on the metathorax in Group A, 
not on that plate in Group B except in one species. Again un- 
described species j diflters in having pro-6 as strong as, and 
slightly longer than, No. 5, and having two or more branches ; 
while its meso-6 is on a plate separate from that of 7. 

No. 7. — In Group A similar to 1 to 5 on the prothorax; 
thick single spine, varying in length according to species on 
the mesothorax; thicker, single, but may be longer or shorter 
than meso-7 on the metathorax. This is, in Group B, stellate 
on the prothorax; single, thick spine (though rather small and 
often branched in two new species) on the mesothorax; thicker, 
more or less curved toward the apex and usually with one or 
two shorter branches on the metathorax; the powelli complex 
often has three shorter branches on the metathoracic spine. 

No. 8. — In Group A short with two or three relatively thick 
branches, or a fairly large tuft of six or more branches on the 
prothorax; invariably a tiny tuft on the meso- and metatho- 
races. In Group B stellate, as No. 1 on the prothorax, also stel- 
late but smaller and with fewer branches on the meso- and 

No. 9. — -In Group A long, stout, single, frayed on all segments, 
except in these species which have this modified into very thick 
spine on one or more segments. In Group B long, branched, 

64 The Philippine Journal of Science ^^*^ 

frayed on all segments, except in new species m which has 
only its meta-9 branched, while its pro-9 is modified into a 
slender spine. 

No. 10. — In Group A modified into very thick spine on the pro- 
and meso-thoraces, and in certain species also on the meta- 
thorax ; but in two species long, single, similar to 9. In Group 
B this hair is more slender, single, either about as long as 9 
or much shorter. The powelli complex, like new species m, has 
this reduced into a slender spine. This unusual peculiarity of 
the powelli complex has not been mentioned before by any of 
the workers who dealt with powelli. 

No. 11. — Tiny, single or branched; comparatively larger and 
with more branches (when branches are present) in Group B 
than in Group A. 

No. 12. — Single, usually longer than 10 on the pro- and meso- 
thoraces in both groups, but that on the metathorax much less 
in rank. 

No. IS. — Most internal of the ventral hairs, variable. In 
Group A either fairly strong though short, single or branched, 
or a tiny tuft on the prothorax; tiny tuft with a few or many 
branches on the mesothorax (apparently absent in two spe- 
cies) ; very thick, branched spine on the metathorax of two 
species ; fairly strong with a few to several branches in others. 
Stellate tuft on all segments of Group B, usually the largest 
on the thorax, particularly that on the metathorax. But un- 
described species j is unusual since its meso-13 is a long, single 

No. H. — In Group A, nearly always similar and equal in 
rank to No. 13 on the prothorax; tiny tuft with a few to 
many branches on the mesothorax; absent on the metathorax. 
In Group B this is stellate on the pro- and meso-thoraces, often 
larger than 13 on the pro-, lesser in rank on the meso-thorax; 
absent on the metathorax. 


Arrangement of the dorsal abdominal hairs is closely similar 
to that of anophelines; the ventral differs slightly. While No. 
10 is ventrally located in Anopheles, this is lateral, anterior 
to 8 in Tripteroides. 

The principal abdominal hairs of Group A, that is, Nos. 1, 
5, 6, 9 and 13 of I to VII, and 7 of I and II are usually either 
thick, spinelike or long, stout, but single. Nos. 1, 5 and 9 are 

78*1 Baisas and Pagayon: Philippine Mosquitoes 65 

stellate or nearly so in three species but these are not of the 
same type as the stellate tufts of Group B. Hairs 0, 1, 5, 8, 
9, 13 and 14 are large stellate tufts in Group B. 

Undescribed species e is unique because its abdominal hairs 
6, 7, and 9, sometimes also 8, arise from a large common plate 
on segments I to V. 

Marked changes in sizes or make-up of certain hairs on dif- 
ferent segments occur in a few species of Group A. Generally, 
however, most of the principal hairs gradually decrease in size 
and length from the anterior to the posterior segments. This 
is equally true, though to a lesser degree, with the large stel- 
late tufts in Group B. 

The marked differences in certain hairs of Group A may be 
used to subdivide the species of the group into smaller units 

(A) Hair No. 1 

(a) About equal in rank, though relatively short on ab- 

dominal segments I-VII . . . undescribed species 6, c, 
and e. 

(b) Very long on IV to VII, short or different on I-III or 

on II and III . . . undescribed species a, d and g. 

(c) Long on all segments . . , microcala and nepenthicola. 

(B) Hair No. 9 

(a) A strong tuft with about ten to more than twenty 

branches on I to IV or I to V .. . undescribed species d 
and g, and nepenthicola, 

(b) Thick, single spine . . . undescribed species c and e, 

(c) Moderately thick, split into two or more branches . . . 

undescribed species a and b, and microcala, 

(C) Hair No. 13. 

(a) Very thick but short, single or divided into two or more 

unequal thick points on I to IV or I to V .. . un- 
described species c and e, 

(b) Being moderately strong, at least on segment I, branched 

. . . undescribed species a, 6, d and g, and microcala^ 


Hair 0. — Present on I or VII. Invariably tiny, sometimes 
wanting on some segments of Anopheles^ but well developed, 
either thick or spinelike, or large stellate tuft in the majority 
of Tripteroides. It is, however, w^eak, though not as tiny as 
in anophelines, in undescribed species c, d, and e of Group A. 
All species of Group B have hair large and stellate. 

6961 5 

66 The Philippine Journal of Science ^^^^ 

Hair 1. — Present on I to VIII. Variable in Group A, ac- 
cording to species, as well as on different segments of certain 
species. Large stellate tuft in Group B except in undescribed 
species j where this is rather weak compared with other tufts. 

Hair 2. — Absent from some segments or some individuals of 
all species in Group A. Stellate, large, present on II to VII in 
Group B. Absent also in what appears to be an unusual kind 
of Rachisoura — newly found, and the only one yet known in the 

Hair S. — On I to VIII. Tiny, single or branched; or a tiny 
tuft in both groups; slightly more developed in some species, 
or on certain segments than in others. 

Hair 4- — On I to VII. Similar to 3, but often longer, in 
both groups. 

Hair 5. — On I to VIII. Spinelike in most species of Group 
A ; long like 6 in others ; strong tuft in one ; small tuft on I to 
III but long hair on IV to VIII in one species. Stellate in 
Group B. 

Hair 6. — On I to VIII. Very long, single on I to VI in Group 

A, much reduced on VII, tiny tuft on VIII. Variable in Group 

B, but long on I to VII in most species; long only on I to VI 
in undescribed species m and n. With at least one of those on 
I to III split into two equal branches, rarely three. One or 
more of those on I to V are branched in some species, while 
some individuals have one or more of those on VI and VII 
split into branches. 

Hair 7. — On I and II nearly as long as 6 with which it arises 
from a common plate; single in all species of Group A, some- 
times branched into two or three in some species of Group B. 
Very much reduced and on separate plate of other segments, 
but relatively long in undescribed species j. 

Hair 8. — On I to VII. Situated on lateral side of segment, 
anterior to 6 and posterior to 10. Tiny, single, or branched 
in Group A; much longer, stellate in Group B. 

Hair 9. — On I to VIII. More or less similar to 5 according 
to species in Group A; invariably stellate like 1 to 5 in Group 
B. When this is stellate in Group A, the branches are mark- 
edly unequal in lengths, quite stout, each tapering apprecia- 
bly from about middle to apex, and terminating into a coarse, 
jagged or clean-cut point. The branches of the large stel- 
late tufts in Group B are not so unequal in lengths, hardly 
taper toward the apices, each ending in a single, double, or 

'«'* Baisas and Pagayon: Philippine Mosquitoes 67 

triple points. Moreover, the branches are evenly spread (when 
m the natural position) into a complete circle, or semi-circle; 
unlike those of Group A which are spread less than 150^ at most' 
Hair 10.— On II to VII. Tiny, single or branched in both 
groups. Situated on the lateral side of the segment anterior 
to 8. 

Hair 11.— On I to VII. Tiny, single, but longer than 10; 
that on VII often branched in both Groups A and B. 

Hair 12.— On II to VII. Variable in position on different 
segments, usually anterior to 11 on II, external to 11 and 
close to the base of either hair 6 or 8 on other segments. Often 
single on the more anterior segments, and branched on the 
others. About the same in either group, except in undescribed 
species j of Group B in which this hair is unusually long. 

Hair IS.— On I to VIII. The most anterior of the ventral 
hairs on segment I, being closer to the thorax than the most 
anterior dorsal hair, 0; but becoming more and more pos- 
terior in position on each succeeding segment in Group A; rel- 
atively constant in location on different segments in Group B. 
Very variable in type in Group A as discussed above; large 
stellate tuft in Group B, being the largest hair on some seg- 

Hair H. — Not found in any species of Group A, or in the 
only known local Rachisoura. Large stellate tuft in Group 
R; the members of the pair very close to each other, except 
on VII, where they are widely apart ; the pair arises very close 
to the posterior border of each segment. 

Abdominal segment VIII bears only five pairs of hairs, the 
homology of which is extremely difficult to determine. More 
for convenience than from any definite reasons, these hairs are 
proposed to be regarded (starting from the most dorsal pair) 
Nos. 1, 5, 3, 6, and 13. No. 1 or the most dorsal pair is usually 
the largest on this segment, but it is a tiny tuft in undescribed 
species d and e. No. 5 is the second most dorsal, ranks usually 
second to No. 1 but longer and more stout where No. 1 is 
weak. No. 3 is usually a tiny tuft. No. 6 equals 5 in most 
species. No. 13 is on the underside of the segment below the 
most ventral comb tooth in Group A, but is on the lateral side 
immediately posterior to the more ventral comb teeth in Group 
B. This is strikingly different in undescribed species j where 
it is the largest hair (tuft) on VIII. 

The hairs on the siphon and anal segment, like the comb-, pec- 
ten-, and saddle-teeth, show group and specific differences, but 

68 The Philippine Journal of Science ^^^^ 

these will be dealt with in subsequent papers on classification 
of Tripteroides. 


1. A complete new set of numeral hair designations for the 
pupa is proposed. This is based on the homology or apparent 
homology between the hairs of the pupa and those of the larva, 
such homology being fairly well indicated when the pupa is 
still in its original position inside the larval skin. The number 
proposed for each pupal seta is exactly the same as that of 
the larval hair to which it corresponds. The widely accepted 
numeral scheme for the hairs of anopheline larva is used and 
referred to here. 

2. A discussion on the elimination, reduction, or increase in 
sizes, and changes in position of hairs during pupation is given. 
It is shown that only three of the nineteen pairs of hairs on 
the head of a Tripteroides larva, and only nine of the forty-two 
pairs on the thorax are represented on the cepalothorax of the 
pupa. According to this, we consider the postocular hairs as 
the only ones that remain of the hairs on the head of the larva ; 
the anterothoracic as the representatives of the prothoracic 
larval hairs; the dorsal and supra-alar as the only two not 
eliminated from the mesothoracic larval hairs; while the meta- 
thoracic larval hairs are represented by the three metanotal 

3. It is also shown that though the great majority of hairs un- 
dergo considerable reduction in sizes during pupation, some, like 
the superior postocular hair and the lateral tufts on abdominal 
segments VII and VIII, increase greatly in sizes. It is like- 
wise indicated that a hair may still exist in the pupal stage even 
when already eliminated from the larval stage; just as a hair 
may not be represented in the pupal stage even when well 
developed in the larva. 

4. Similarly, it is shown that the general similarities in ar- 
rangement of pupal hairs between different genera, subgenera, 
groups and species cannot, in some cases, be a guide to identify 
the different setse. This is due to the varying positions of cer- 
tain hairs in relation to each other. Moreover, the relative 
positions of some hairs are altered when the pupa is freed from 
the larval skin, and stretches to assume its normal shape. Ab- 
dominal hairs 3, 4, 6, and 8 are specially affected by the stretch- 
ing of the pupa. 

'^' 1 Baisas and Pagayon: Philippine Mosquitoes 69 

5. Differences in hairs on the head between the larva of an 
Anopheles and that of a Tripteroides are discussed and shown 
to be more in types and arrangement rather than in actual 
number. Only the post-clypeal hair or No. 4, and that which 
springs near the apex of the maxillary palp or No. 16 are not 
represented in Tripteroides. Changes in position of hairs are 
well indicated by the inner clypeal or No. 2 of Group B, sub- 
genus Tripteroides. In two species it is located on the fronto- 
clypeus, but in all the others, including those of Group A, this 
springs from the pre-clypeal plate. Movement or instability 
in position of this hair is further indicated by individuals which 
have a member of the inner clypeal pair located on the pre- 
clypeal plate while the other member arises from the fronto- 
clypeal plate. 

6. The usual method of regarding the most anterior of the 
frontal hairs in culicines as B seems erroneous for Tripteroides 
where this is not only the most anterior but also the most in- 
ternal. In numeral designation this is No. 5 and should, there- 
fore, be C instead of B, The hairs on the head are discussed 

7. The number and arrangement of hairs on the thorax and 
abdominal segments I to VII closely resemble those of anophe- 
lines. The differences displayed on segments VIII to X are 
obviously due to the differences in the development of these parts 
between an Anopheles and a Tripteroides. The different tho- 
racic and abdominal hairs are discussed individually. 


Plate 1 

Fig. 1. r. nitidoventer. Right half, dorsal side of abdominal segments IV 
and V, showing larval hairs in solid lines, and pupal seise in 
situ (dotted lines), a and b = tips of branches of the stellate 

2. T. microcala. Left half, dorsal side of abdominal segments V, 

VI and VII, showing larval hairs in solid lines, and pupal se- 
tae in situ (dotted lines). 

3. T. (undescribe) species d. Right half, dorsal side, abdominal seg- 

ment I of larva, showing larval hairs in solid lines, and pupal 
setae in situ (dotted lines), 

4. r. monetifera. Ventral side right half of head showing larval 

hairs in solid lines and pupal setae in situ (dotted lines). Note 
the increase in size of pupal hair No. 14. 

5. T. monetifera. Right half, dorsal side of the thorax, showing the 

larval hairs in solid lines, and the pupal seise in situ (dotted 

6. Dorsal portions of two larval heads showing difference in posi- 

tion of hairs 2 and 5. 

(a) left side, undescribed species a of Group A. 

(6) right side — Tripteroides nitidoventer of Group B. 

Note. — The diffrent figures are not drawn in the same scale. 

Plate 2 

Fig. 1. Reproduced from Plate VII of Baisas, showing pupal hair notations 
slightly modified from the scheme of authors. Represents dorsal 
side, left half, of a Culex (Culex), 
2. Reproduced from Plate IV of Rozeboom and Knight, showing the 
numerical pupal hair notations they introduced. Represents 
Anopheles farauti, 

Plate 3 

Fig. 1. T. dyari of Group B. Parts of a pupa showing the metanotal 
and abdominal hairs. Right half represents the dorsal side; 
left half, the ventral. 

2. T. (undescribed) species m. Anterior portion of larva showing 

right half, dorsal hairs; left, half, ventral hairs. The dotted 
lines represent the visible pupal setae. Insertion at left of head 
is an enlargement of part of pre- and fronto-cypeal plates. 

3. T. larva (undescribed) species 6. Terminal part of abdomen 

(lateral view). ct = comb teeth; pt = pecien teeth; st = saddle 

4. Thorax of Tripteroides larva, undescribed species b, (Left half 

represents ventral side, right half dorsal side). Note shortness 
of all hairs. 


72 The Philippine Journal of Science 

Plate 4 

Fig. 1. Comb teeth, saddle teeth and pecten teeth of undescribed species m, 

2. Head of Tripteroides larva, undescribed species 6. (Left half ven- 

tral side, right half, dorsal side). 

3. Tripteroides undescribed species m. Posterior portion of larva 

showing right half, dorsal hairs; left half, ventral hairs. Seg- 
ments VIII-X from lateral aspect. Dotted lines represent visi- 
ble pupal setae. 

4. Abdominal segment IV of Tripteroides larva, undescribed species 

(Left half represents ventral side; right half, dorsal side). 
Note. — shortness of all hairs, st = saddle teeth. 

Baisas: Pup^ and Larvje of Tripteroides.] 

[Philip. Jour. Sdi., tS, Nd. 1. 


Baisas: Pupje and Larvae of Tripteroides.] 

[Philip. Jour. Sci., 78, No. 1. 


Baisas: Tupje and Larv^ of Tripteroides.] 

[Philip. Jour. Sci., 78, No. 1. 


Baisas: Pup^ and Larv^ of Tripteroides.] 

[Philip. Jour. Sci., 78, No. 1. 





By Ernesto M. Paras 

Of the Chemical Section, Culion Leper Colony^ 

Bureau of Hospitals, Department of Health, Philippnes 

The study of calcium in leprosy, which in the past several 
years had attracted the attention of many leprosy investigators, 
has been centered chiefly on the question of whether or not in 
those afflicted with this disease a calcium deficiency exists. The 
interest in this study undoubtedly had its inception from the 
clinically manifested evidence of the occurrence of bone salts 
resorption in the hands and feet of the afflicted, especially in 
those of the advanced stage. The approach into the problem 
has been directed mainly to the determination of calcium bal- 
ance and the total concentration of calcium in the whole blood 
or serum. However, as will be observed in the review given 
below, such methods only yielded variable results in the hands 
of the different workers. Further study of this important prob- 
lem is deemed necessary. 


Underbill, Honej and Rogert^ reported that two leper sub- 
jects by the balance method, especially the more advanced, 
manifested a definite tendency to retain calcium. They in- 
terpreted this finding to indicate that in the organism a de- 
ficiency of this element exists. Along the same line, Boulay 
and Leger^ obtained results on three cases which failed to 
confirm this observation of the aforementioned workers. They 
reported that the retention tendency was observed only in one 
and in the other two there was hyperexcretion. From the 
examination of eleven cases, Reyevski* reported the balance 
as positive but only in those whose leprosy remained stationary. 
This result, including that which he obtained from blood exam- 
ination, suggested to him that ''during the acute stage of the 

' Published with the approval of the Director of Hospitals. 
2Underhill, F. F., J. A., Honej, and L. J. Bogert. Jour. Exp. Med., 
S2, (1920), 41-63. 

3Bouley, A., and M. Leger, Sec. Path. Exot., 15, (1932), 865. 
4Reyevski, A. S. Arch. f. Schiff-u Trop. Hyg., 34, (1930), 651-57. 


74 The Philippine Journal of Science ^^^ 

disease decalcification takes place, blood [Ca]++ falls and 
[K]+ rises." 

In the examination for total calcium content in whole blood, 
Concepcion and Salcedo^ reported their findings on 37 lepers 
as normal except in a few paroled cases where the calcium 
content showed a slight abnormal increase. Leman, Liles and 
Johansen,^ from the serum examination of 54 cases, failed 
to observe a deviation from normal in the serum content of 
this constituent. In similar examination of 47 cases, Wooley 
and Ross'' reported this also to be normal. Villela,^ on the 
other hand, from the serum analysis of 113 cases, claimed 
that 51 or about 45 per cent gave values that were abnor- 
mally low. He reported this abnormality to occur mostly in 
the neural type and not in those with lepra reaction. The 
study on 70 lepers by Cruz, Lara and Paras ^ however, 
showed that it is in the great majority of those with reaction 
where the value was observed below normal. Only in the 
non-reacting did they find it to remain within normal range. 
These findings appear to have gained support from the works 
of Badenoch and Byron ^^ on 54 cases, and of Herrera ^^ on 62 
cases. In a study by Nishikawa ^^ on 38 cases the serum level 
was reported to be slightly lower than normal. Otsuka and 
Yoshinaga" reported it normal in the macular form and low 
in nodular, especially in the neural type. 


To state briefly, the universally accepted concept today con- 
siders calcium as it exists in this body fluid to be constituted 
mainly of the diffusible and the non-diffusible form. The for- 

5 Concepcion, I., and I. Salcedo. Jour. Phil. Is. Med. Assoc, 6 (1926) 154. 

6 Leman, I. I., R. T. Liles, and F. S. Johansen. Am. Jour. Trop. Med., 
Baltimore, 7 (1927) 61. 

7 Wooley, J. S., and H. Eoss. U. S. Pub. Health Rpts., j^6 (1931) 641. 

8 Villela, G. G. Int. Jour. Leprosy, 6 (1938) 63. 

9 Cruz, M., C. B. Lara, and E. M. Paras. Jour. Phil. Is. Med. Assoc, 
8 (1928) 216-221. 

10 Badenoch, A. G., and F. E. Byron. Trans. Roy. Soc Trop. Med. Hyg., 
26 (1932) 253. 

11 Herrera, R. M. Siglo Med., 95 (1925) 394. 

12 Nishikawa, T. La Lepre S (1932) No. 2. 

13 Otsuka, R., and T. Yoshinaga. Trans. 10th Meet. Jap. Lep. Assoc, 
La Lepre, 9 (1938) Suppl. 32. 

14 For comprehensive review of various works on the subject, readers 
are referred to Schmidt, C. L. A., and D. M. Greenberg. Physiol. Review, 
51 (1935) 297. 

^^' 1 Paras: Blood-Chemistry Studies in Leprosy HI 75 

mer is regarded as practically all in an ionized state and is 
the fraction of physiologic and clinical importance, whereas the 
latter is looked upon as nearly all bound to proteins and be- 
lieved to be biologically inert. These two fractions are said 
to account for most, if not all, of the concentration of total 
calcium in this body fluid. 

It can be seen from this concept that the above-cited studies 
based on the estimation of total calcium content in blood are 
evidently devoid of clinical significance unless the protein con- 
centration is also known. Further approach to the problem 
on the basis of this newer knowledge would thus appear to 
be of no little interest. 


The examination of the diffusible and the non-diffusible cal- 
cium simultaneously with the albumin and globulin in the 
serum of lepers was attempted only by Wooley and Ross. "^ 
From a study of 53 lepers, they reported the average for the 
diffusible and the albumin-globulin ratio as considerably low, 
increasing abnormally in the non-diffusible and the globulin. 
Only in a few instances did they find the level approximating 
the value for the control. Thus, they were led to the conclusion 
that **the clinically improving cases are accompanied with an 
increase in the diffusible and the albumin-globulin ratio and a 
decrease in the percentage of the globulin and the non-diffusible 

The evidence obtained from the above study, particularly with 
regard to the diffusible, undoubtedly is significant and it is 
deemed necessary to have this field studied farther. It is with 
the ultimate objective to help supply the much needed informa- 
tion that the present work has been undertaken. 


From our previous studies of the inmates of this Colony 
it was our observation that cases with complication of lepra 
reaction, especially the febrile type, are most readily suscep- 
tible to undergo abnormal changes in the concentration of 
their serum constituents. Accordingly, in the belief that they 
would make ideal subjects to meet the present objective, they 
were taken to constitute most of the cases. Thus, of the 19 
selected, 12 were from this group, 3 were those without reaction 
but advanced, and 4 were also without reaction, but quiescent. 


The Philippine Journal of Science 



Blood of about 20 cc. was drawn from each patient for the 
analysis of various constituents. After having allowed the 
drawn blood to clot, the resulting sera were worked out at 
once. A portion of the serum was used for the determination 
of the total calcium, albumin and globulin and the rest was 
subjected to ultrafiltration for the analysis of the diffusible and 
the non-diffusible calcium. The procedure for ultrafiltration 
and the colorimetric analysis for albumin and globulin were 
carried out in detail according to the improved technique de- 
scribed by Greenberg and Gunther ^^. The ultrafiltrate was 
analyzed for the diffusible according to the gasometric method 
of Van Slyke and Sendroy ^^ Only those ultrafiltrate giving a 
negative biuret test were used. For the analysis of total cal- 
cium, the Tisdall modification ^^ of the Kramer-Tisdall method 
was employed. The value for the non-diffusible was obtained 
by substracting the value obtained directly for the diffusible 
from that of total calcium. 


The individual findings obtained in lepers are given in 
Table 1. 

Table 1. — Results obtained in 19 cases of leprosy, showing partitions of 
serum calcium and proteins. 

Milligrams per 100 cc. serum 

Grams per 100 cc. 






Condition of leprosy 












With lepra reaction. 




















































































Clinically negative. 




































19-. __ 







15 Greenberg, D. M. and L. Gunther. Jour, Biol. Chem., 85 (1920-30) 491. 
Greenberg, D. M. Jour. Biol. Chem., 82 (1929) 545. 

16 Van Slyke, D. D., and J. Sendroy. Proc. Sec. Exp. Biol. Med., 23 
(1926) 167. 

17 Tisdall, F. F. Jour. Biol. Chem., 56 (1923) 439. 


Paras: Blood-Chemistry Studies in Leprosy HI q^j 

To facilitate comparison, the maximum and minimum limits 
and the average value taken for the control and those obtained 
for the three groups of leprous cases are arranged in Table 2. 

Table 2.- 

-Figiires from Table 1 compared with control arranged to show 
only the minimum^ maximum and the average. 

Const! tuenta 


Reaction cases 







Total calcium 


3*. 77 
















3*. 34 

Diffusible calcium _ 

Nondiffusible calcium 


Globulin __ 

Quiescent cases 

Advanced cases 







Total calcium . 



















Diffusible calcium 

Nondiffusible calcium _ 


Globulin _. 

Total calcium. — An examination of the data compiled in the 
said tables will show that the serum level for total calcium 
suffers abnormal reduction only in lepra reaction group. This 
finds confirmation in the previous study of this constituent 
by us 1^ Of the 12 cases in this group, 7 or about 58 per cent 
show values that are definitely below normal. No appreciable 
difference in the average value can be noted between the dif- 
ferent groups except in that with reaction where it is observed 
to be definitely below normal. 

Diffusible calcium, — The serum level of this constituent, both 
in the reaction group and in the quiescent and the advanced, 
is, without exception, within the range of normal limits. Com- 
pared with the control, the average (Table 2) appears higher 
in the 3 groups of cases, the advanced having the highest, 
next the reaction group, and least the quiescent. 

Non-diffusible calcium, — In the case of this constituent, the 
serum level in 10 out of 12 reaction cases (about 83 per cent) 
is subnormal, and only in 1 out of 3 cases (about 33 per cent) 
in the advanced group where it is found to be abnormally 
low. In the quiescent group, it is all well within normal 
range. The average, compared with the control, is practically 

78 The Philippine Journal of Science ^^ 

the same as that in the quiescent and appreciably low both in 
the advanced and the reaction groups. 

Serum proteins. — The fate for the albumin in general ap- 
pears to follow the same trend as in the case for total cal- 
cium, that is, abnormal reduction only in cases with lepra 
reaction complication, otherwise normal. Of the 12 cases in 
this group, 10 or about 83 per cent, show values that are definite- 
ly subnormal. The average in this group is markedly low, but 
in the other two groups it does not appear to differ appreciably 
from the normal. In the case of the globulin, its serum level 
is raised markedly above normal in 9 of 12 cases (75 per cent) 
in the reaction group and practically normal in the quiescent 
and advanced groups. The average value is definitely above 
normal in the group with reaction and practically normal in the 
other two groups. 


Before discussing the results just presented it may be well to 
touch first upon those evidences which have given rise to the 
contention of the existence of a deficiency in calcium in this 
disease. These are: {a) the finding of Underhill and his asso- 
ciates,2 based on calcium balance measurement, of a definite 
tendency of the organism to retain this element, with the ap- 
parent support it gained from the work of Reyevski;^ (&) 
the abnormal reduction in the serum total calcium level ^^ ^' ^^^ ^^' 
12.13. and (c) the lov/ finding in the serum diffusible.'^ As 
regards the first, it should be realized that the various data 
reported in this study, besides the fact that they lack com- 
plete accord and are in conflict with that obtained by Boulay 
and Leger, ^ are only based on two cases. It should further be 
reckoned that the application of the balance method to the 
investigation of human subjects, especially in diseased condi- 
tions, could not likely yield decisive results, even if it is carried 
out with the most scrupulous care, because of the complicating 
factors involved. The situation in the case of that revealed 
from total calcium examination is still less convincing because 
on the basis of the newer knowledge, the claims cited, besides 
being also variable and quite in conflict with those of others,^'^*'^ 
could not have clinical meaning unless the serum protein con- 
centration is also known. 

Coming now to the diffusible, the present study, as seen 
from the results just presented, failed to confirm the low 

78, 1 Paras: Blood-Chemistry Studies in Leprosy HI 79 

finding reported by Wooley and Ross for this constituent. 
As already pointed out, not a case of the nineteen patients 
subjected to the analysis, irrespective of the presence of major 
complication like lepra reaction and the advancement of the 
disease itself, was its level found to have undergone abnormal 
lowering. But this disagreement, as will be noted, is more ap- 
parent than real and in fact their finding for this constituent 
can serve, instead, to establish the results obtained in this work. 
Thus, if on re-examining the data of the aforementioned 
workers the low normal limit of 4.2 for the diffusible is used 
instead of the 5.5 which the said workers took for this purpose, 
43 or 81 per cent of the 53 cases whom they subjected to 
the examination will be found to have values that fall well 
within the range of normal, and only in 10 or 19 per cent 
where this drops below normal. But even in these cases, 
5 will be seen to approximate closely the limit of normal and 
only 3 would be in the zone which, according to the exper- 
ience of other investigators, could only be compatible with the 
existence of tetany that is caused by parathyroid insufficiency. 
Whether the marked abnormality in these latter cases is caused 
by the existence of an actual calcium deficiency or due to 
faulty methods in the analysis, this paper will not attempt to 
discuss. It may seem pertinent, however, to mention that 
in the absence of tetany, low values could result from a poorly 
prepared membrane that has been utilized in the ultrafiltration or 
compensation dialysis and from a membrane that has been badly 
impregnated through repeated use. Which of the two limits 
just considered could justifiably represent the actual or at least 
approaching the true value is of course of importance to deter- 
mine. For this matter, we can only state for the present that 
we find justification in the adoption of 4.2 from the consider- 
ation of the following: (a) our previous survey on serum total 
calcium and serum proteins of those residing in this Colony 
yielded values which revealed to be quite comparable with those 
reported in the literature, thus showing that climatic factor, 
etc., do not have any appreciable influence, if at all, upon the 
concentration of these constituents. It does not seem likely, 
therefore, that the diffusible low level would also deviate ap- 
preciably, if at all, from this value; (&) it represented a much 
larger series of different groups of normal individuals than that 
in the value taken by Wooley and Ross. In the latter the nor- 
mal individuals that it represented can be said to be composed 
of practically one class of such individuals whose standard of 

go The Philippine Journal of Science ^^^^ 

living, etc. are above average; (c) it is the widely accepted 
value among investigators; and (d) it is in accord with that 
obtained from any dialysis experiments by various investi- 
gators and from that obtained by the use of direct biologic 

It could be seen plainly from the above that there was no 
valid reasons to consider this serum constituent to be defi- 
cient in this disease as Wooley and Ross w^ould lead us to be- 
lieve. Theoretical considerations of the mechanism that is in- 
volved in the equilibrium relationship between calcium and 
proteins, which the recent studies of McLean and Hastings ^^ 
have proved to exist in this body fluid, would seem to provide 
means by which this contention could find further support. To 
facilitate discussion on this point let us for the moment touch 
briefly upon this mechanism. From the works of the above- 
cited investigators it appears that the calcium ions, protein ions 
and the calcium salts of proteins, behave as if there was equilib- 
rium between them; that this behavior appeared to obey mass 
action law and could be expressed by the equation 

tCa]"^"^ X [Protein]' 
[Ca Proteinate] 

K (1) 

in which K varies with the pH of the normal serum. Now, on 
the basis of the conditions by which the constituents under con- 
sideration in this study are known to exist, the application of 
the above equation would justify limiting the determining fac- 
tors for the equilibrium, mainly, to the diffusible calcium, the 
non-diffusible calcium and the albumin. The equation, accord- 
ingly, could evidently be represented by 

[Diffusible Ca]"^"^ X [Albumin] 

[Non-diffusible Ca] 
( Ca-albuminate) 

= K (2) 

As may be seen then from this latter equation,!^ in calcium de- 
ficiency, mass action law would have us expect the level of both 
the diffusible and the non-diffusible to come down low. But as 

18 McLean, F. C, and A. B. Hastings. Jour. Biol. Chem., 108 (1935) 

19 The present data could not be treated strictly quantitatively, in the 
first place because they are not expressed in molecular concentrations 
which is a requirement of the expression, and secondly, they are believed 
to approximate only the conditions. Necessarily, therefore, they can at 
best be discussed qualitatively and this is believed sufficient to meet the 
purpose of this study. 

^«' ^ Paras: Blood-Chemistry Studies in Leprosy III 31 

the results show, while the non-diffusible actually suffered ab- 
normal reduction in accordance with this expectation, the dif- 
fusible level failed to have undergone similar fate. It can thus 
be seen that the contention in the existence of such a deficiency 
can neither, on this basis, be satisfactorily explained. Neither 
could the data of Wooley and Ross be, because, even granting 
for the sake of argument, that their finding for the diffusible 
was really low, tne non-diffusible, which should also be low as 
expected to be found in conditions where there is such deficiency 
was shown, instead, to be increased abnormally. 

Now, to view the results from another angle, equation 2 can 
be seen to help explain satisfactorily the fall in the level 
of the albumin as the one accountable for the reduction that 
occurs in the non-diffusible. The results obtained in this study 
appears to help affirm this assumption. Assuming this to 
be the case it becomes quite plain that the low results obtained 
in the total calcium could be due to the fate that befell the 
non-diffusible, the diffusible having shown to have not under- 
gone abnormal lowering. As for the increase in the globulin, it 
may be surmised that this probably, considering the well-known 
functional value of this constituent in the body defense mecha- 
nism, is a product elaborated at the expense of the albumin in 
an effort to overcome the ill-effects caused by the lepra reaction 
attack and such other causes. 

As can be noted from the discussion the reference to the 
equation was practically limited in explaining the deficiencies. 
While it may not be entirely out of the scope of the present 
work, it is thought worthwhile considering also the fact that 
from the equations given, by increasing the total concentra- 
tion of protein, the reaction trend would be toward increas- 
ing the concentration of the constituents that suffered deficien- 
cies and this tendency will continue to be so with increase of 
supply of the required substance until the dissociation constant 
of the equilibrium has been restored to its original state. It 
thus appears quite suggestive that protein therapy or adequate 
protein diet, or both, may prove beneficial in the treatment 
of lepra reaction cases where the above-cited deficiencies are 
found to occur mostly. It may be pointed out that calcium 
drug may have similar effect in the matter of restoring the 
equilibrium dissociation constant by elevating the level of the 
non-diffusible and may therefore also prove beneficial. As a 

6961 6 

82 The Philippine Journal of Science ^^^ 

matter of fact the experience ^o in this Colony has shown this 
to have really appreciable ameliorating effect in the treatment 
of the reacting cases. However, as already discussed above, 
the focus of disturbance is not in the calcium but in the pro- 
tein component system and for this fact protein may prove 
to be more effective. 


The present investigation involving analysis of the serum 
for its content of total calcium, diffusible calcium, non-diffusible 
calcium, albumin and globulin in nineteen lepers has been un- 
dertaken with a view to help solve the long controversial ques- 
tion of whether or not calcium deficiency exists in leprosy. 

The inadequacy of the data compiled from previous studies 
of the earlier investigators based on the determination of cal- 
cium balance and of the concentration of total calcium in the 
whole blood or serum was discussed. 

In this study the presence of major intercurrent diseases, like 
lepra reaction and the advancement of the disease itself, was 
shown to have no appreciable influence, if at all, upon the nor- 
mal level of the diffusible. The low results observed by Wooley 
and Ross for this constituent were shown to be more apparent 
than real and that they rather serve, instead, to establish the 
finding reported in this study. 

The great majority of the cases with lepra reaction revealed a 
general abnormal lowering in the serum level of total calcium, 
non-diffusible calcium and the albumin and an abnormal shift 
above normal of the globulin. The non-reacting cases failed to 
show appreciable abnormal alterations, if at all, in these con- 
stituents. The explanation for the conditions found in the 
reaction was given on the basis of an equilibrium relation- 
ship assumed to exist in this body fluid between calcium and 

From the consideration of the facts given in the above dis- 
cussions, the writer is led to conclude that in leprosy there is 
no deficiency of calcium, at least in so far as the serum phys- 
iologic calcium that is measurable in terms of the diffusible 
is concerned. The abnormal rise in the level of the globulin 

20 Roxas-Pineda, E., C. Nicolas and C. B. Lara. Jour. Phil. Is. Med. 
Assoc. (1928), 207. 

78,1 Paras: Blood-Chemistry Studies in Leprosy III 83 

is believed to be at the expense of the albumin which causes its 
fall to below normal and this effect upon the latter in turn 
causes the lowering in the non-diffusible and in the total 

Protein therapy is suggested as worth giving a trial in the 
treatment of leprosy cases suffering from lepra reaction at- 
tack, especially in those manifesting low albumin in the serum. 


By Wm. S. Hamm and Arthur Avery 
Of the Philippine Fishery Program, U, S. Fish and Wildlife Service 

Salt production through the solar process has, perhaps, been 
practiced for centuries in the Philippines and has, through 
the years, gone through considerable evolution towards more 
efficient methods in many of the producing provinces. The 
methods now in use, however, are not as efficient nor are 
they as productive of a pure salt as the solar salt producing 
methods practiced in some other parts of the world. 

Cox et al (1915), Ablan et al (1940), Montalban (1930), 
Seale (1914), and Herre et al (1920) all refer to the fact 
that the poor quality of the locally produced salt has a dele- 
terious effect on the fish preserved with it. Tressler (1920) 
has shown that calcium, magnesium and sulfates as impurities 
in salt retard the penetration of salt into fish during curing. 
Dunn (1946) has shown that herring fillets are struck through 
sooner during brining if the salt is pure. Hess (1942) has 
shown that impurities in salt affect the keeping quality of cod 
press juice. Shelf life of salt fish may, therefore, be extended 
by the use of pure salt. Tressler (1923) and Phalen (1917) 
point out that salt for many industrial purposes must be of 
high purity. Table 1 gives the results of some analyses of 
local and foreign salts. This table illustrates that the local 
salt is not as pure as solar salt produced in other areas and sug- 
gests that salt of equal quality could be produced here if some 
changes were made in the present practices. 

Table 1. — Analyses of samples 

of sea salt 


Rican 1 

California 2 

Bahaman « 


Sand and insolubles 

Per cent 








Per cent 

Per cent 

Per cent 

Sulfate fSOO 


Calcium (Ca) _ _ 


Magn^ium (Mg) _ _ _ 


Sodium chloride (01) calculated _ 


Chloride (CI). 

Ferric oxide and alumina (Fe203 and AI263) _ 

* Produced by a modified improved Oriental process. This salt had not been washed. 
2 From Tressler, A. K. B\2reau of Fish Doc. 884 (1920). 


86 The Philippine Journal of Science ^^*^ 

Local methods. — In the solar salt process the first require- 
ments are a hot sun, preferably brisk breezes of low humidity, 
and large areas of impermeable land near the sea. These re- 
quirements are found in several provinces in the Philippines 
but the industry is for the most part located in Rizal, Bulacan, 
Cavite, Bataan, and Batangas provinces. In these provinces, 
the so-called, improved Oriental process is practiced. In this 
process, as employed in the Philippines, areas used as fish 
ponds in the wet season are used to produce salt during the 
dry months. 

Most of the fish pond area is employed for concentration 
ponds where the seawater, 10 to 20 centimeters deep, is evap- 
orated down to a more or less saturated solution. Usually, 
for ease in operation, this is done in three or more separate 
ponds. The raw seawater may be evaporated in the first pond 
from 3.5° Be to 4.5° Be, in the second to 6° Be, in the third to 
8°Be, and in the fourth to about 12 to 15°Be. In the concen- 
trating ponds, the sediment settles, and the iron sulfide, cal- 
cium carbonate and some of the calcium sulfate are deposited. 
This brine, at about 15° Be, is then dipped with buckets into 
crystallizing ponds where it is evaporated almost to dryness 
and the salt recovered. These latter ponds are about 5 me- 
ters by 7 meters equipped with board sides and tile bottoms, 
and usually represent from 1/10 to 1/15 of the entire area at 
the salt works. 

Everyday, during the season, the ponds are filled to a depth 
of about 1 cm. and late in the afternoon after each day of 
evaporation the salt is raked into a pile on the side of the 
crystallization bed and the pond refilled. This process is contin- 
ued throughout the season, without any change, until the brine 
level builds up with a heavy brine containing a large percentage 
of the magnesium salts. When this happens, the salt producer 
will probably let the pond go for a day or more without adding 
fresh brine, and will then, after harvesting the salt, splash out 
of the pond the remainder of the brine. Owing to the high 
magnesium content, this method of handling produces a most 
inferior product. It is even possible at times to find crystals of 
pure magnesium sulfate intermixed with the salt. As explained 
above, it is the common practice of the salt producers in the 
Manila area to fill the crystallization beds with brine of 12 to 
17° Be which is the point at which calcium sulfate begins to 
precipitate, and thus practically all the calcium sulfate is found 

'^' 1 Hamm and Avery: Philippine Solar Salt Industry 87 

in the final product as an impurity. Further, it may be noted 
that the practice of using an unsaturated brine in the crystal- 
hzers results in a lower yield since a portion of each day 
IS spent in evaporating the brine to a point where salt will 
begin to form at 25° Be. 

A great part of the salt produced is sold as it is harvested 
but a considerable portion is stored, both by speculators and 
producers, in the hope that there will be a better market dur- 
ing the non-producing season. During the period of storage 
up to twenty per cent of the weight of the salt is lost owing 
to the moisture being absorbed from the air in such quantities 
that the piles of salt, literally, drip. This loss could be les- 
sened by producing a purer salt which would have a lower 
percentage of the highly hygroscopic magnesium and calcium 

In the provinces of Ilocos Norte, Ilocos Sur, La Union, and 
Pangasinan, salt is prepared by the old Oriental leaching 
process. Their process is described by Tressler (1920). Ablan 
et al (1940) in an extensive field study of this process came to 
the general conclusion that this method of salt production 
was not economic and suggested, that in certain places, the 
improved Oriental process might be profitably introduced. 

California practice. — The American salt making process in 
general is similar to the improved Oriental process but the 
details of the process vary greatly. The operations are con- 
ducted in various ponds. These are: (1) the storage or in- 
take pond into which the salt water is received from the bay, 
(2) concentrating ponds, and (3) crystallizing ponds. 

Some of the works have automatic gates which open when 
the tide is at flood and close when it begins to ebb. The salt 
water remains in the intake pond for various lengths of time, 
and is pumped into the concentrating ponds to a depth of 25 
to 50 cm. before the next sea water is taken in. It then 
goes through the different ponds, becoming more concentrated, 
until it reaches the crystallizing ponds. It is run into these, 
when it reaches a strength of 25° Be, to a depth of 15 centi- 
meters or more. This depth of brine causes the deposition of 
large crystals which, over the extended period of evaporation, 
grow into massive solid crystals of nearly pure salt. Here the 
brine evaporates until it has reached a density of 29° Be, at 
which time the bittern is either discarded or evaporated in 
other ponds to a density of 32°Be. This salt, formed from 


The Philippine Journal of Science 


the heavier bittern, is less pure. The process is repeated un- 
til the end of the season or until a thick enough bed of salt 
is obtained to make the harvesting economical. 

The sea water is usually pumped from the intake pond to 
the concentrating ponds, through which it flows by gravity. 
The brine may then flow by gravity or be pumped into the 
crystalizing ponds, depending on the lay-out of the land. 

All of the ponds have mud bottoms and often a split is made 
in the harvesting. That is, all the salt except the bottom 5 
centimeters will be taken first and then the bottom part will 
be removed and used where a cheaper salt can serve. In the 
modern plants, machinery is now used to remove the salt from 
the beds. Often the harvested salt is washed with brine from 
the concentration ponds and with fresh water just after it is 
harvested. By this treatment some of the producers get salt 
of over 99.5 per cent sodium chloride content. It is possible 
to get a relatively pure product with this simple washing proc- 
ess because the crystals are large and solid. Impurities are 
only on the outside surfaces of the crystals and the mother liq- 
uor and washwaters drain free. Further, in the storage of 
the large solid crystals under conditions of high humidity, less 
salt is lost than when storing small crystals. 

Experimental results. — Several experiments were conducted 
on a salt farm in Las Pinas, Rizal. The first experiment was 
designed to determine the effect on purity of salt obtained 
from brines of different densities. A bed 5 meters by 7 meters 
was filled to capacity, about 5 cm. in depth. No salt came 
down until the fourth day and on the 5th day rather large 
hollow hopper shaped crystals formed. Table 2 gives the 
analyses for magnesium in the various samples. 

Table 2. — Effect of brine density on magnesium content 

Days after filling 

Brine density 


Per cent 


Per cent 


5 - 




8 - 

Sample from control pond 

This table illustrates how a somewhat greater purity can be 
obtained at the lower brine densities, but this reduction in 

^^' ^ Hamm and Avery: Philippine Solar Salt Industry gg 

magnesium content is of too small a magnitude to be of any 
practical significance. This is because the standard practice 
under local production methods involving daily harvests results 
in the formation of relatively small hollow salt crystals. These 
crystals have a relatively high surface to volume ratio which 
retains a great deal of the impure mother liquor. Further they 
are of such shape that the mother liquor does not drain as 
freely as it would from large solid crystals. Simple washing 
experiments on these salts also failed because the purity could 
not be improved enough to make the washing practical. 

The failure of the foregoing experiment in developing 
a method of producing a purer salt led to two series of 
experiments designed to illustrate how a greater operational 
efficiency could be achieved. In these experiments one crys- 
talizing pond was filled with the brine at about 14 ° Be— the 
density at which it is usually added to the crystallizing pond. 
After it had evaporated to about 25° Be it was pumped into a 
second pond. In these experiments over a two week period an 
average daily production of 56 kilograms was realized from the 
experimental plot, whereas only 38 kilograms were harvested 
from the control plot. The control plot was operated exactly 
the same as the regular procedure for this particular salt works. 
This experiment indicates that for a given total area of salt 
works it should be possible to reduce the area of crystal- 
lization beds by about one third the present space devoted to 
such use and still obtain the same yield. The area then 
made available can be used as additional concentration 
ponds to concentrate the brine to 25° Be. This should result 
in considerable savings to the operators as the investment in 
preparing crystallizing ponds is quite high. It should also 
reduce harvesting costs to some extent. 

Conditions affecting rate of evaporation, — It is apparently the 
belief of the salt producers that evaporation is more rapid 
from shallow ponds, and also that evaporation proceeds more 
rapidly over the tile in the crystallization ponds than from deeper 
ponds with a mud bottom. To determine any bases of these 
contentions experiments were made in small pans. Clear white 
enamel photographic pans and black plastic photographic pans 
were used. The tile was set in mud placed in an enamel pan. 
In the experiment conducted over a mud bottom an enamel pan 
was used with about 2.5 cm. of soil. The measurement of evapo- 
ration was determined by weighing after eight hours in the sun, 


The Philippine Journal of Science 


and again after standing overnight, 
of these experiments. 

Table 3 gives the results 

Table 3. — Effect of bottom surface and depth on evaporation. 

Type of pan and bottom 

Tiie bottom. 

Mud bottom 

Black plastic 

Clear white enamel 



Black plasti* 


Initial depth 

Kg. of water 

of water 

evap. per 

sq. m. 

8:00 a.m. to 


U:00 p.m. 

















Kg. of water 
evap. per 

sq. m. 

U hrs. 


These experiments show that the rate of evaporation is not 
greatly influenced by depth of water at the depths tested when 
the bottom does not reflect the sun's rays. It is indicated, 
though, that when the bottom reflects the sun's rays the evap- 
oration proceeds more rapidly from the deeper water. The 
difference in the rates of evaporation between the experiments 
with tile and mud bottoms is not considered to be significant. 
There is, obviously at the depths tested, an advantage in hav- 
ing the bottom of a color which will absorb the heat rays from 
the sun. The difference in the evaporation for 24 hours be- 
tween the deep and shallow water levels for the black plastic 
containers can be explained on the basis of the greater latent 
heat of the deeper brine at sunset. 

Table 4 gives temperature and density variations at various 
points on a salt farm over the period of most active evapora- 
tion. The depths in the various ponds were about 2 cm. in 
the crystallizing pond and about 4 cm. in the concentration 
pond and at least 15 cm. in the ditch. It is an interesting 
observation to note that the temperature of the brine in the 
ditch was higher at the bottom than at the surface. This is 
explained by the fact that the brine in the ditch evaporates 
throughout the day and is then lower in level than the crystal- 
lizing ponds when refilled. When additional brine from the last 
concentrating pond at a lower density is brought in it does not 
mix with the remainder of the heavy brine in the ditch but 
just flows in over the surface. The sun then heats the mud 
bottom of the ditch and this heat is only slowly absorbed by 
the brine. Actual tests showed that the brine at the bottom 
of the ditch was of greater density than that at the surface. 

78.1 Hamm and Avery: Philippine Solar Salt Industry 91 

Table 4. — Temperature and density value at various places on a 

salt farm 3 

Ditch bottom 

Ditch surface 

Crystallization pond 

Concentration pond* 























. 39 



































































3 Temperature is in degrees centigrade and density in degrees Baume. 
* The concentration pond here referred to was a small tile-surfaced pond, normally used 
for crystallization. 

Table 5 gives the evaporation and rainfall during the salt 
making season for the San Francisco Bay area and also the 
corresponding data for the Manila area. 

Table 5. — Evaporation and rainfall in millimeters 


December. - 


February- _ . 








Salt Season total - 

San Francisco 








20. U 


Net evap. 














Net evap, 



From these data it is seen that there is about 16 per cent 
greater net evaporation in the Manila area than in the San 
Francisco area during the salt making season. We may there- 
fore assume on the basis of the weather alone that local con- 
ditions are satisfactory for the introduction of the California 
method. The following disadvantages may be considered, how- 
ever, in the introduction of the new system: 

1. The use of deep brine in all the ponds might make it nec- 
essary to use pumps. Lack of electrical power and cost of 
pumping equipment might make this type of operation too 
expensive. . 

2. The problem of displaced labor might be acute in certain 

producing areas. 

92 The Philippine Journal of Science ^^^ 


1. The operating efficiency can be increased and the cost of 
production somewhat decreased by concentrating the brine to 
the point at which salt begins to form, prior to placing brine 
in the crystallization beds. 

2. Favorable conditions exist for the introduction of more 
efficient production technics. 

3. In order to produce a purer salt the present practices 
must be considerably modified. 

4. Because of the crystal characteristics of Philippine salt, 
it is difficult to purify it by washing. 

5. Within certain limits the depth of the evaporating brine 
has no significant effect on the rate of evaporation. 


1. Serious consideration should be given to the application 
of the California method of salt production in the construction 
of new salts works. 

2. Old operators should review their production costs and 
consider the application of more modern methods. 

3. Additional experiments should be made demonstrating the 
feasibility of improved methods under the prevailing conditions. 


The authors wish to thank Dr. Santos of Paraiiaque, Rizal, 
who made his salt works available for these experiments and 
demonstrations. Appreciation is also extended to the Philippine 
Bureau of Fisheries for their assistance on this project. 


Ablan, L., Jose R. Montilla and Basilio M. Martin. Salt making 

industry of northern Luzon. Philip. Jour. Sci. 72 (1940) 319. 
. Cox, Alvin J. and T. Dar Juan. Salt industries and resources of the 

Philippine Islands. Philip. Jour. Sci. 10 (1915) 375-93. 
Dunn, J. A. Paper presented before annual meeting of the Institute of 

Food Technologist March 18, 1946. 
Hess, Ernest. Effects of various salts on preservation. Jour. Fish. Res. 

Bd. Can. 7 (1) (1942). 
Herre, a. W. and Jose Mendoza. Bangos culture in the Philippine Islands. 

Philip. Jour. Sci. S8 (1929) 450-509. 

■^8' 1 Hamm and Avery: Philippine Solar Salt Industry 93 

Mgntalban, Heraclio R. Investigations on fish preservation conducted 

at Estancia, Panay, P. I. Ill Philip. Jour. Sci. 42 (1930) 809-333. 
Phalen, W. C. Technology of salt making in the United States Bureau 

of Mines. U. S. Bureau of Mines Bulletin 146 (1914). 
Seale, Alvin. Preservation of commercial fish and fishery products in the 

tropics. Philip. Jour. Sci. 9 (1914) 1-6. 
Tressler, D. K. U. S. Bureau of Fisheries. Document 884 (1920). 
Tressler, D. K. Marine products of commerce. The Chemical Catalog Co, 

Inc. (1923) 762 pp. 



Of Iloilo College, University of the Philippines 


Scylla serrata (Forskal), S. oceanica (Dana), S. tranqueba- 
rica (Fabricius), and the new variety S. serrata var. parama- 
mosain comprise the group of crabs that are called alimango 
(Tagalog and Visayan), rasa (Ilocano), and alama (Pangasi- 
nan). They are generally regarded as our most valuable 
crabs, and it is rather strange that we know very little about 
their biology. The present work has for its aim the following 

(1) To study the biological peculiarities of the animals, so 
that measures for their propagation can be adopted. 

(2) To study the mode of reproduction and the development 
of the young to the size at which sexual maturity is attained. 
" (3) To study their bionomics— the economic relations of the 
different phases in the life history of the crabs. 

(4) To show the advisability of enacting wise fishery laws 
designed for their conservation and proper exploitation. 

The work started with the assumption of previous authors 
that S. serrata comprises the only representative of the genus 
in the Philippines; hence, the external morphological features 
were not scrutinized very closely. The writer, however, was 
aware from the beginning of the existence of what apparently 
was a variety, which fisherman around Manila Bay call buhk, 
meaning motley-spotted. These crabs are characterized by 
their carapace, abdomen (in female only), and legs bemg 
bizarrely ornamented with rather large polygonal pigmented 
areas having shades of grayish to whitish, the spots bordered 
by purplish or dark-brown lines. The color markings are 
more distinct in medium-sized crabs, but in the bigger and older 
ones the pigmented areas on the carapace become mdistmct, 
owing to their being obscure d by deposition of sediments or 

^This paper proposes the revision of the genus to include three species 
and one new variety based on external morphology and |ametogenesis^ 
More detailed data on spermatogenesis and oogenesis of the different forms 
will be treated in a subsequent paper. ^^ 

96 The Philippine Journal of Science ^^^^ 

algal growth which give the carapace a muddy appearance. 
The general coloration is greenish with shades of brownish gray 
in some places of the body including the chelipeds and legs. 
The bulik crabs grow bigger than any of the other forms, some 
specimens measuring about tw^enty-three centimeters across the 
greatest width of the carapace. 

In some places in Quezon Province the bulik group is known 
as banhatvin, and the other kinds mamosain. For conve- 
nience, the two terms are adopted throughout the discussion 
for grouping the crabs. The mamosain group is characterized 
by the absence of color markings like those found among the 
banhawin, or bulik. Some whitish spots may be present on 
the carapace and legs, but, generally, they are indistinct and 
are mostly confined on the last legs. The general coloration in 
7namosain is something like a combination of rust-color (ferru- 
gineous) and dark brown; the shades of color may vary, but 
the general greenish tint, characteristic in banhawin, does not 
appear. The mamosain (referring only to S. serrata) as dis- 
tinctly hole-dwellers; hence, they are commonly met with in 
brackish swamps. Among the fishermen in the Batangas Prov- 
ince, they are known as alimango sa butas (crabs in holes) ; 
this habit, is in contrast to the decided propensity of the 
banhatvin toward a roving life. 

At first there was no plan to go into the question of tax- 
onomy. But in the course of the study the following problems 
occurred : 

(1) Variations in the modes of spermiogenesis. 

(2) Somewhat corresponding deviations in the process of 

(3) Differences in the form and structure of chromosomes. 

These features together with the external differences dis- 
cussed above as well as differences in bionomics led the writer 
to suspect the existence of more than one species. 

The crabs used in these studies were bought from the mar- 
kets in Baguio, Manila, Batangas, Batangas Province, and from 
places around Dagatdagatan Experimental Station at Malabon 
in Rizal Province. 


Crabs of different sizes were dissected during different 
months — ^March, April, May, June, July, August, September, 

^^'* Estampador: Revision of the Genus 97 

and December. The conditions of the gonads were studied 
to determine the probable range of the spawning season, and 
the relative sizes at which these crabs attain sexual maturity. 
Pieces of the testes and of the ovary obtained by cutting 
across the organs were fixed, using different fixatives, such 
as Zenker; Carnoy and Lebrun; modified Beuin with urea; 
modified Flemming, including Benda: chrome-aceto-osmic : Ga- 
tenby: strong Flemming without acetic acid, and Flemming's 
strong solution with 40 per cent formaldehyde in place of 
acetic acid. The best result in chromosome fixation was ob- 
tained with modified Beuin of the following formula: 

Picric acid, saturated solution parts.... 75 

Formaline (strong) do 15 

Glacial acetic acid do 10 

Urea crystals gr 0.5 

With Gatenby's-Flemming's strong fluid without acetic acid, 
and iron hsematoxylin, good results were obtained showing Golgi 
bodies and fat globules. 

Different combinations of stains were tried. The best re- 
sults were obtained with iron hsematoxylin, acid fuchsin and 
anilin blue. This combination stains the chromatin black, the 
cytoplasm and fibers red in various shades, and the metaplasm 
of the secretory cells, blue. 

Double embedding in collodion and paraffin was employed 
for pieces of ovary containing mature eggs. After dehydra- 
tion the tissues were soaked in collodion dissolved in equal 
parts of ether and absolute alcohol for about two weeks; then 
passed through either cedar oil or clove oil, in the meanwhile 
the collodion film formed removed around the tissues. The 
tissues were then passed through equal parts of absolute al- 
cohol and benzene; through pure benzene, and then benzene 
and soft paraffin (in solution). The tissues stayed in the last 
solution about a week. The tissues were then embedded in 
pure hard paraffin. Good sections of eggs were obtained by 
this method. The usual method of preliminary infiltration with 
soft paraffin in the oven was avoided to reduce the chances of 
the tissues becoming too brittle from overheating. The sec- 
tions of ovary were cut 7 microns thick; those of the testes, 
5 microns thick. 


To the late Dr. Hilario A. Roxas, formerly head of the Depart- 
ment of Zoology, University of the Philippines, I wish to express 

6961 7 

98 The Philippine Journal of Science ^®*^ 

my gratitude for his kindness in extending to me the facilities 
at the Dagatdagatan Experimental Station. Doctor Roxas was 
at the time the chief of the Division of Fisheries, Department 
of Agriculture and Commerce. I am also indebted to Dr. Deo- 
gracias Villadolid, now Director, Bureau of Fisheries, for ex- 
tending to me the same privileges. My thanks are also due to 
Mr. Heraclio Montalban for his ready cooperation in placing at 
my disposal the necessary conveniences. I am also grateful to 
Mr. Felix Arriola and to other members of the station staff 
who helped me in securing the material from fishermen. And 
lastly, to Dr. Leopoldo S. Clemente, the present head of the 
Department of Zoology, University of the Philippines. I wish 
to take this opportunity to express my appreciation for his 
patience in going critically over the histological portion; and 
to Dr. Leopoldo B. Uichanco, dean of the College of Agriculture, 
University of the Philippines, and to Dr. Marcos A. Tubangui, 
of the Institute of Hygiene, for their valuable suggestions and 


(Swimming Crabs) 

Portunidas MiERS, Challenger Report, Zool. 17 (1886) 169. 

Carapace depressed, moderately transverse, usually widest 
at last antero-lateral spines. Front horizontal. Orbits and 
eye-stalks nearly always of moderate length. Lateral teeth 
usually from five to nine. Last legs usually adapted for swim- 
ming, with terminal joints ovate, flatly expanded. 

Genus SCYLLA de Haan, 1833 

Scylla De Haan, Fauna Japonica, Crust (1833) p. 11; A. Milne- 
Edwards, Ann. Sci. Nat. Hist. Zool. (4) 14 (1860) 228, 249; Ar- 
chiv. du Mus. 10 (1861) 347. 

Carapace transverse, bread, moderately convex, with an al- 
most unbroken surface. Front proper well delimited from in- 
ner supra-orbital-angles, cut into four teeth; its breadth (ex- 
cluding supra-orbital aijgles) between a fourth and fifth of 
greatest breadth of carapace. 

Antero-lateral borders oblique, arched, longer than postero- 
lateral, cut into 9 teeth of nearly equal size. 

Orbits without any dorsal inclination; two nearly closed fis- 
sures in the upper wall ; inner angle of lower border dentiform 
and prominent. Antennules folded nearly transversely. Ba- 

^^'^ Estampador: Revision of the Genus 99 

sal antennal joint produced to form a lobule lying in the orbit; 
flagellum, of good length, standing in the orbital hiatus. 

Epistome sufficiently long fore and aft, not sunken. Buccal 
cavern squarish, broader than long; efferent branchial chan- 
nels cavernous, but not defined by ridges. 

Chelipeds massive, longer than any of legs; arm wrist and 
hand with definitely placed spines; hands deep and full, not 
prismatic, not costate. 

Legs stout, moderately compressed; in fourth pairs merus 
and carpus shortened and broadened, and propodite and dac- 
tylus typically foliaceous for swimming. 

Abdomen of male rather broadly triangular, consisting of 
5 segments, third to fifth terga being fused. First tergum 
much concealed beneath carapace. 

SCYLLA SERRATA (Forskal). Plate 1, fig. 1 

Cancer serratus Forskal, Descript. Anim. p. 90. 

Lupea tranquebarica H. Milne-Edwards, Hist. Nat. Crust. 1 (1834) 

Scylla tranquebarica Dana, U. S. Explor. Exped. Crust. 1 (1852) 

270; Stimpson, Acad. Nat. Sci. Philad. (1858) 38; Stimpson, 

Smith Misc. Coll. 49 (1907) 75. 
Scylla serrata De Haan, Fauna Japonica Crust. (1850) 44. 
Scylla serrata De Hann Fauna Japonica, Crust. (1860) 44; Krauss, 

Sudofr. Crust, p. 26; Miers, Ann. Mat. Nat. Hist. 5 (1880) 238; 

Miers, Chall. Kept. Brachyura 17 (1886) 185; De Man, Arch. 

fur Naturges., Jahr. 53 (1888) 332; Chia-Jui Shen, The Crabs 

of Hongkong. The Hongkong Naturalist 3, No. 1 (1932) 32. 

Carapace about two-thirds, or a little less, as long as broad, 
practically smooth, except for a faint granular ridge running 
obliquely inwards across either branchial region from last spine 
of anterolateral border. 

Front cut into four lobes or bluntish teeth of about equal 
size and prominence. Antero-lateral border cut into nine 
sharply acuminate teeth of about equal size. Posterior border 
forming a curve with posterolateral borders, points of junction 
sometimes slightly thickened. 

Merus of external maxillipeds oblique but not having antero- 
external angle distinctly produced in a lateral direction. 

Chelipeds not quite twice length of carapace in adult male, 
but shorter than in female and young male. Arm with three 
spines on anterior border, and two on posterior border one ter- 
minal, other submedian; a strong spine at inner angle of wrist, 
outer angle being rounded and armed with one, or sometimes. 

100 The Philippine Journal of Science ^^^ 

two spines or teeth. Hand with three spines or tubercles, one 
being in front of apex of wrist joint, other two being side by 
side behind finger- joint, outer of these two sometimes obso- 

Legs unarmed: dactyli strongly sulcated, their anterior and 
posterior margins fringed with brushlike hairs. 

Abdomen of male broadly triangular: oval in young fami- 
lies, but becomes more circular in outline in big females. 

Remarks. — Carcinologists have hitherto recognized only one 
local species under this genus. Miers (1886) recognized only 
S. serrata; and Stimpson (1907) seems to be of the opinion 
that S. serrata and S. tranquebarica are synonyms. Both Da- 
na (1852) and Stimpson placed S. oceanica as a variety under 
S. tranquebarica. The present study attempts to establish three 
species and one new variety. 

De Haan, in adopting Scylla for the generic name, must have 
had in mind the habitat and habits of the crabs. Scylla (Lat. 
from Gr. Skylia) is described in Greek mythology as a sea 
monster, dwelling in a cave in a precipitous cliff that guarded 
the entrance to a passage. She is said to devour any prey 
that comes within her reach. This allusion does not fit the 
banhawin (bulik) crabs, because they do not live inside the 
holes. The allusion fits S. serrata. Kraus (in Sudafr. Crust.) 
relates: "this species lives in great deep holes, and wears the 
dingy earthy color of its residence. They sit at the opening 
of their holes when the tide is coming in, to snap at the food 
which it brings them." 

In Scylla serrata the outer of the two spines behind the 
fingerjoint is described as obsolescent (Plate 1, fig. lb. This 
is so in the true mamosain, where in some cases especially in 
big males, the spine is reduced to a mere inconspicuous ves- 
tige. Among the banhawin group both spines are conspic- 
uous ; the outer one, however, is generally smaller. 

Between the true banhawin and the true mamosain, there 
are two groups of individuals that seemingly occupy the in- 
termediate positions in general coloration, one group closely 
approximating the banhawin, while the other resembling close- 
ly the mamosain. On the whole the differences in the shades 
of color and in the extent of pigmented areas can be passed 
on easily as simple variations that can be attributed to such 
factors as age and environment. But when these disimilar- 
ities are supplemented by (a) differences in form and size 
of the chromosomes, and (6) differences in the process of 

^^»^ Estampador: Revision of the Genus 101 

spermiogenesis and ookinesis, they then assume greater signifi- 
cance. Obviously, they are due to fundamentally deep-seated 
factors inherent in the distinctive make-up of the individuals. 
To serve as bases for description and for convenience, the 
group that closely resembles the true banhawin is designated 
as parabanhawin and those that closely approximate the true 
mamosain (S. serrata), as paramamosain. The true banhawin 
is identified in this paper as S. oceanica, and the parabanhawin 
as S, tranquebarica, the paramamosain is regarded as a variety 
under S. serrata. 

SCYLLA OCEANICA (Dana) Banhawin. Plate 1, fig. 2. 

Scylla tranquegarica var. oceanica Dana, U. S. Explor. Exped. 
Crust. 1 (1852) Stimpson, Smith. Misc. Collection 49 (1907) 75. 

Diagnosis. — This species can be distinguished from S. serrata 
by the following characteristics : 

1. The general coloration is predominantly greenish, the 
shades in some places of the body becoming grayish green in 
bigger specimens. The carapace, legs, and abdomen are or- 
namented by large pigmented areas of whitish gray bordered 
by purplish dark-brown lines. These ornamental markings are 
absent in S. serrata in which only irregularly disposed small 
whitish spots may be seen on the carapace and on the last legs, 
and the general coloration is usually deep f errugineous brown ; 
the shades of color may sometimes vary, assuming a light 
purplish brown. 

2. S. oceanica ordinarily grows bigger. It is not unusual to 
see specimens that measure about nine inches across the widest 
portion of the carapace. Probably bigger specimens may exist. 

3. Brushlike setae (Plate 2, fig. 2) are present quite abun- 
dantly over the carapace of S. oceanica. While in S. serrata 
the setae appear to be shorter and are apparently confined only 
to the hepatic region just below the orbit. 

4. They differ in habitat: S. serrata referred to in this paper 
as the mamosain, usually lives in holes; while S. oceanica (the 
banhawin) seemingly prefers a nomadic life, and is commonly 
caught in open seas. 

5. Relatively, more pores perforate the carapace of S. ser- 
rata. The hypodermal papillary tubes (Plate 2, fig. 3) pro-, 
trade through these pores. These minute tubes are extensions 
of the hypodermis and are, obviously, respiratory in function. 
(See section in bionomics.) 

102 The Philippine Journal of Science *^^ 

Other differences as revealed by cytology: 

6. Their chromosomes differ in their physical make-up and 
in form. 

7. The modes of spermiogenesis and ookinesis also differ. 
These and other cytological differences will be brought out 
more in detail in the course of the discussion. 

Remarks. — Both Dana (1852) and Stimpson (1907) con- 
sider S. oceanica as a variety under S. tranquebarica. Stimp- 
son pointed out the existence of two sufficiently distinct var- 
ieties. He gave as the distinguishing characteristics of S. oce- 
anica the following: (1) the frontal teeth are blunt, with the 
median incision deepest; (2) the posterior teeth of the antero- 
lateral margin is longer than in the other. Stimpson described 
the coloration in the species as follows: "carapax olive green 
above, white below ; hands often reddish and spotted with green 
exteriorly; fingers always red, feet pale greenish, with dark 
spots below." 

H. Milne-Edwards (1834) characterized Lupea tranquebarica 
as the biggest species of Portunidse, and, according to him, they 
may attain the length of from six to eight inches; the color is 
grayish green, and they inhabit the Asiatic seas. Presumably 
his specimen was S. oceanica, since this is the variety that 
grows very big, and the big ones are, apparently, sea-dwellers. 

Notwithstanding their close similarity with regards to gen- 
eral coloration, the writer proposes to raise S. oceanica to the 
category of a distinct species and to include therein forms 
which are: 

(1) Predominantly greenish or grayish green on carapace 
and legs, including chelipeds; these parts ornamented by large 
pigmented areas. 

(2) Those lengths of chelipeds (in male) not more than twice 
the length of carapace; hands not reddish or purplish as a 
whole, though splashes of reddish color may appear. 

If the writer were to consider one as a variety under the 
other, he would choose S. tranquebarica, on the ground that 
the general coloration and color markings on this group seem 
to be more variable; and on the basis that the germ cells of 
S. tranquebarica are heterogametic, they exhibit sexual diga- 
mety and dimegaly. While those of S. oceanica are apparently 
homogametic, a condition which implies more stability of 

'*'^ Estampador: Revision of the Genus 103 

SCYLLA TRANQUEBARICA (Fabricius) Dana. (Farabanhawin). Plate 3, fig. 1. 
Pertunus serratus Ruppel, Krabben die rothen Meeres, p. 19, pi. 2, 

fig. 1. 
Lupea tranquebarica Milne-Edwards, Hist. Nat. Des Crust. 1 (1834) 

Scylla serrata De Haan, Fauna Japonica, Crust. (1850) 44; Krauss, 

Sudafr. Crust., p. 25. 
Seylla tranquebarica Dana, U. S. Explor. Exped. Crust. 1 (1852) 
270, pL XVI, fig. 6; Stimpson, Acad. Nat. Sci. Philad. (1858) 
38; Stimpson, Smith. Misc. Collection 49 (1907) 75. 

Diagnosis. — Differs from S. oceanica by the following charac- 

1. General coloration varying from deep purplish drab- 
green to lighter in shade; overall greenish background which 
is characteristic of the true banhawin {S. oceanica) generally 
showing only on the carapace and on some portions of the 

2. Large pigmented areas generally distinct only on the last 
legs; indistinct on the other legs and on abdomen of female. 
Chelipeds usually purplish; fingers red, often tipped with 

3. Relative differences in the size of the chelipeds more ap- 
parent, especially so in males, where the chelipeds are enor- 
mously big in relation to the body. In adult males the length 
of the chelipeds is more than twice that of the body. 

4. As in S. oceanica, the nurse cells in the gonads are dis- 
tinctly vacuolated; in size the auxocytes compare favorably with 
those of the aforementioned. 

5. Unlike those of S. oceanica, however, the sperms of S. tran^ 
quebarica exhibit sexual digamity and the ova show sexual di- 

megaly. . 

Remarks.— The variations in general coloration and in color 
markings within the species are quite diverse; it is difficult 
to determine whether they should be regarded as simple ex- 
ternal variations, or something that preclude existence of races 
or varieties. These need further cytological studies. 

According to Stimpson, these species are found on the shores 
of the continent of Asia, chiefly in muddy estuaries. They 
are sold in great numbers in the markets of Hongkong and 

Canton. - 

In the Philippines the market supply of these crabs gener- 
ally comes from the fishponds; considerable quantities of the 
crabs are also caught from the muddy shores of bays, coves, 
and chiefly the muddy estuaries. 

104 The Philippine Journal of Science *^^ 

SCYLLA SERRATA var. Paramamosain var. nov. Plate 3, fig. 2. 

Characters. — The general configuration of the body is very- 
similar to that of the typical species. In both the body is 
relatively thicker, appears to be more compact, and the ca- 
rapace looks more convex than that of either S. oceanica or 
S. tranquebarica. The H-like mark on the carapace is rela- 
tively faint; the figure is deeply impressed in both S. oceanica 
and S. tranquebarica. 

Variety paramamosain differs from the typical species by 
the following characters: 

1. Median pair of frontal lobes slightly more anteriorly 
produced than the two laterals. 

2. Outer of the two spines at the base of the fingers, though 
smaller than the inner, neither obsolescent nor dentiform; it 
is quite prominent. In the species, especially in big males, the 
outer spine becomes vestigial. 

Some cytological differences 

3. The ova of paramosain exhibit sexual dimegaly. The con- 
dition in the sperm has not been determined. 

4. The form and the apparent physical constitution of the 
chromosomes of paramamosain differ from those of the ma- 
mosain (S. sererata). 

5. The modes of spermiogenesis and ookinesis also differ in 
these two groups. 

Color. — It is rather hard to distinguish offhand the variety 
from the typical species in so far as general coloration is con- 
cerned. There is an apparent tendency in the variety, how- 
ever, to assume a lighter shade of color, which is usually 
brownish with splashes of gray. Usually, there are patches 
of bluish green at the angles of the joints of the legs and che- 
lipeds. The spines in the chelipeds are of this color excepting 
the tips. Small pigmented areas, but rather irregularly dis- 
posed, may be seen on the last legs; these areas consist of 
groups of whitish spots, and as in the species, they are not bor- 
dered by distinct lines. 

On the whole the shades in color and the color markings may 
vary with such factors as age or environment. Also, there are 
no distinctive external differences. Hence, despite differen- 
ces in the physical constitution of their chromosomes and 
in the modes of spermiogenesis and ookinesis, it seems safer 
with the present state of our knowledge tentatively to place 
paramamosain as a variety. 

^^'* Estampador: Revision of the Geniis 105 

Key to the Philippine Scylla. 

aK General ground color greenish, grayish green or olive-green; some- 
times with shades of purplish or reddish generally on chelipeds and 
legs. Contour of body not very distinctly convex; H-like mark on 
carapace relatively deep. 
b \ Carapace and legs, including chelipeds, predominantly greenish or 
grayish green; large polygonal pigmented areas present on all 
legs, including chelipeds, and on abdomen in females; chelipeds in 
adult males not more than twice length of carapace — ^S. oceanica, 
b ^ Carapace olive-green, purplish with greenish tint, or purplish brown; 
large polygonal pigmented areas distinct only on last legs, and 
a few may be seen on the abdomen in females; chelipeds often 
reddish or purplish, length in adult n^les more than twice length 

of carapace S* tranquebarica, 

a ^. General ground color deep rusty-brown, purplish brown to light brown- 
ish gray; dorsal contour of body more distinctly convex; H-like 
figure on carapace relatively less deep. 
b\ Color varying from deep rusty brown to the purplish brown; outer 
spine on base of fingers obsolescent, becoming vestigial on big- 
ger specimens; frontal teeth about same level S. serrata. 

b ^ Color generally brownish gray with patches of bluish green at joints 
of legs and chelipeds; median pair of frontal teeth slightly pro- 
duced anteriorly; outer spine on base of fingers unequal but not 
obsolescent S. serrata var. paramamosain var. nov. 

Dimorphism. — Usually the right cheliped is bigger than the 
left, but in cases which are apparently not very uncommon, 
the left is bigger than the right. The difference in size between 
the chelipeds is more pronounced in males than in females, 
and is quite exaggerated among the paramamosain, but more 
so in S. tranquebarica (parabanhawin) . In these groups, es- 
pecially in males of tranquebarica, the chelipeds appear to be 
enormously big in relation to the size of the body. The hand is 
thickly inflated and smooth; the fingers are very large, thick, 
and strong, curved and furnished with large thick teeth. In 
males the circumference of the hand measured at the base of 
the fingers almost equals the width of the carapace. 

Changes in the form of the abdomen as the crab approaches 
maturity. — There are individuals with abdomen oval, instead 
of triangular, and considerably broader than the abdomen in 
.males, but not broad enough to compare with that of a ma- 
ture female (Plate 2, figs, a, b, and c). These are imma- 
ture females; their sternal concavity is deep as in males, and 
the vulva is rather reduced in size. The abdominal append- 
ages are also relatively reduced in size. Studies made on 
the stained sections of the gonads of two crabs — one, a ma- 
mosain, and the other a banhawin — revealed the presence of 


The Philippine Journal of Science 


oocytes and immature ova in different stages of development. 
In the individuals studied, there were no indications of the 
presence of developing male germs, so that apparently no 
copulation had previously taken place. As the crab grows 
older, the abdomen increases in width, and becomes more or 
less rounded. 

As stated previously, the present study was started with 
the presumption that there is only one species. Hence, in 
the earlier part of the investigation, no attempt was made 
to classify the crabs. The discovery of existing differences, 
which obviously point to the existence of more than one spe- 
cies, made it necessary to change the methods. A systematic 
checking of the results thus far obtained is also made im- 
perative. Some of the materials that were used in this 
checking are included in the Table 1 to show the degree 
of dimorphism as shown in the difference of the size of the 
chelipeds; and also, to show the difference in the general 
coloration and color markings. 

Table 1. — Showing some of the characteristic features of the Philippine 
species of the genus Scylla, 


of carapace 

of palm a 

General coloration and 
color markings 





S. oeeanica {hariJiQwin) : 




























Greenish with large pigmented 
areas on carapace, legs and 





Female _ 

Carapace drab-green due to algal 

Light brownish purple; big pig- 
mented areas on last legs only. 
Crab still soft. 

Carapace greenish; chelipeds pur- 
plish, large pigmented areas on 
last legs only. Circumference 
of chelipeds almost the same 
as width of body. 

Deep rusty brown; scattered 
whitish spots on last legs. 

Deep purplish brown; whitish 
spots on last legs. 

S. tranguebarica (paraban- 


S. serrata (mamosain) : 

Immature female 


S, serrata var. paramoaain: 

Light purplish brown; with 
splashes of bluish green on 
joints of legs and chelipeds, 
whitish spots on last legs. 

Same color marking as foregoing. 

Dark purplish brown; with 
splashes of ferrugineous areas 
on other parts of body. 



« Measurements were taken at the base of fingers. 
*> Missing. 

78»i Estampador: Revision of the Genus 107 


The separation of S. oceanica from S. tranquebarica as dis- 
tinct species is amply justifiable. The external differences 
of these species supported by other characteristic features in- 
dicated by variations in the modes of spermiogenesis and of 
ookinesis; and by the peculiarities in the form and physical 
constitution of their chromosomes, etc. — these are quite 
sufficient bases. 

Aside from their ecological preferences, the banhawin (S. 
oceanica) are distinctly different from the mamosain {S. 
serrata) in their external features. The presence of large 
polygonal pigmented areas that mark the carapace, all of the 
legs including the chelipeds and abdomen in females, make 
the former group easily recognizable from the latter. In ma- 
mosain only whitish spots may be seen here and there, on 
the carapace and on the legs, but on closer examination these 
spots appear to be groups of pigmented dots, and they are 
not delineated by distinct boundary lines. There is no pos- 
sibility of mistaking a mamosain from a banhawin. 

Because of their close similarity in general external appear- 
ance and in coloration, the writer tentatively places parama- 
mosain as a variety under S, serrata, notwithstanding their 
cytological differences. 

Undoubtedly, the study on the genetics of these groups will 
prove very interesting. Experimental breeding will furnish 
evidences from fertility-sterility test held by many as an ob- 
jective method of demonstrating species validity. On the 
whole, capacity to interbreed with fertile offspring is regarded 
as a very significant evidence in any consideration of the 
degree of consanguinity of closely similar stocks. 

The segmented character exhibited by the chromosomes, and 
in some cases the tendency of chromosome-constituents to- 
wards the formation of a somewhat loosely arranged groups, 
are altogether significant. Those arrangements of chromoso- 
mal components make possible the so-called "translocation'', 
or ^'displacement,'' in which a piece of chromosome becomes 
transversely detached and joins the chromosome of another 
pair. This segmental interchange gives rise to a new linkage 
group, since the genes contained in the translocated section are 
now free from the genes with which they were previously as- 
sociated; they may become a part of another linkage group 
constituted by the chromosome to which they have become 

108 The Philippine Journal of Science 

attached. This had been observed in many animals; it may 
have been taking place in these crabs too — a possibility which 
is hinted by the physical constitution of the chromosome. 

Translocation, or displacement, is regarded as having a great 
influence in the evolution of new species, since it provides a 
rapid means of chromosome differentiation which must have 
a great influence on the evolution of new races, varieties, sub- 
species, and species. 

From observations embracing differences in general color- 
ation, color markings, and other external features, the writer 
suspects that besides the ones discussed in this paper, there 
are at least other variations. 

The existence of mamosain {S. serrata) in considerable num- 
ber in certain regions in the Philippines appears to be cor- 
related with the existence of extensive mangrove swamps. The 
relation is obvious, on the ground that these crabs are distinct- 
ly fossorial mud-dwellers. The mangrove with its stiltlike roots 
and a variety of other plants with similar growth habits, espe- 
cially Senneratia pagatpat Blanco, S. caseolaris (L.) Engl., and 
Aegiceras corniculatum (L.) Blanco, are common on the quiet 
coastal areas. The tangle of roots holds debris that were 
brought down by rains and that drifted with the tides until 
it decays. In and upon the mud is found a remarkable as- 
semblage of marine, freshwater, and terrestrial animals. A 
habitat with such conditions gives ample protection to the 
crabs, and wonderfully fit their predaceous and scavenger 


1. Chia-Hui Shen. The Crabs of Hongkong. The Hongkong Naturalist 

S (1932). 

2. De Haan. Fauna Japonica, Crust. (1850) 44. 

3. De Man. Arch, fur Naturges. Jahr. 5S (1889) 332. 

4. EsTAMPADOR, E. P. Checklist of Phil. Crust. Decapods. Philip. Jour. 

Sci. 62 (1937). 

5. MiERS, E. J. Challenger Kept., Brachyura 17 (1886). 

6. Milne-Edwards, H. Hist. Nat. Crust. 1 (1834) 448. 

7. Stimpson. Acad. Nat. Sci. Philad. (1858) 38, Smithsonian Misc. Coll. 

J,9 (1907) 75. 


Plate 1 

Fig. 1. (a). Scylla serrata (Forskal) ; female. 

(6). Scylla serrata (Forskal); left cheliped of male showing the 
spines behind the finger joint; the outer has become vesti- 
2. Scylla oceanica (Dana) ; female. 

Plate 2 

Fig. 1. (a). Abdomen of a female S. oceanica; width of carapace 19.5 
cm.; length, 13.6 cm. 
(6). Abdomen of young female S. oceanica: width of carapace 11.6 

cm.; length, 8.2 cm. 
(c). Abdomen of a female S. serrata var. paramamosain: width of 
carapace 11.6 cm.; length, 8.3 cm. 

2. Terminal portion of a brushlike seta from the carapace of S, 

oceanica (banhawin) ; X 1350. 

3. A hypodermal papillary tube taken from the carapace of the same 

specimen; X 1350. 

Plate 3 

Fig. 1. Scylla tranquebarica (Fabr.) Dana; male. 

2. Scylla serrata var. paramamosain var. nov.; female. (Note the 
two spines behind the finger joints, which are almost equally 



I run. IP. JiH'K. Sn., 7S, No. 1. 

|! f?'Hiflf lWl fffj! | il|l|f|l)f^^ 

/'^- ,*:!Zf'#_ 








iPmi.iF. JoML Hci., 78, No. 1. 

tM^^^s:^i3 4^^ '4 :, s^TW . 

% S .» L U ii ■* >- -h 111 :-4 

w^»t. .f mmmMM 


Estampadob: Stupiks on Stn lla, I. 

I Pmijp. ,}tn'iL 8ci„ TS, No. 1. 


By Gloria D. Manalo and Augustus P. West i 

Manila elemi is one of the well-known minor forest products 
of the Philippines. It is a resin that is contained in the bark 
of the pili tree, Canarium luzonicum (Blume) A. Gray, and 
oozes out when the bark is cut. 

It is used in pharmacy as an ingredient in plasters and oint- 
ments. It is employed in making paints, varnishes, lacquers, 
and also as a constituent in the manufacture of plastics and 
printing inks. Locally it has served as varnish and as ma- 
terial for caulking boats. Forest dwellers roll the resin in 
leaves and use it as an illuminant. A large torch, thus prep- 
ared, gives a light that burns for many hours and can be seen 
far away. 

For many years Manila elemi has been exported from the 
Philippines but the quantity shipped annually has varied con- 
siderably. In 1940 the amount sent abroad was valued at 

By operating a pilot plant for the manufacture of com- 
mercial products from Philippine resins, like Manila elemi and 
copal, very likely important new industries could be developed. 

Resin obtained from different locations usually contains more 
or less foreign matter (leaves, twigs, etc.) and varies in com- 
position. For industrial purposes it is desirable to ascertain the 
quality of Manila elemi and also the amount of individual resin 
constituents. We have devised an accurate and convenient 
method of analysis and the details are given in this paper. 
There are also included the data on some characteristic sam- 
ples of Manila elemi, as determined by this procedure. 

Clover 3 and also Bacon * have carried out extensive resear- 
ches on Manila elemi collected in the Philippines. Their in- 
vestigations were confined particularly to the volatile oil ob- 
tained from this resin. A review of their work^ is given 
in Bulletin 20 of the Philippine Bureau of Forestry. 

1 Formerly of the Bureau of Science, now Institute of Science. 

2 Annual Report. Insular Collector of Customs, Manila. (1940) One 
peso equals 50 cents United States currency. 

3 Philip. Jour. Sci. 2 A (1907) 1. 

4 Philip. Jour. Sci. 4 A (1909) 93. 

6 West, A. P., and W. H. Brown. Philip. Bur. For. Bull. 20 (1920) 40. 


112 The Philippine Joumal of Science ^^*^ 

The name elemi is a term that is applied to a variety of 
resinous products obtained from different countries and hav- 
ing different botanical origins. In the Philippines only one 
elemi resin is collected and this is obtained from Canarium 
luzonicum which grows abundantly at low and medium alti- 
tudes in the primary forests of Luzon, Mindoro, Ticao and 
Masbate. It is the only Philippine tree that gives this resin 
in sufficient quantities for profitable collection. 

In addition to Canarium luzonicum there are various other 
species of Canarium in the Philippines. Canarium ovatum 
grows abundantly in southern Luzon. It produces nuts known 
as pili nuts and is said to give an annual yield of about 33 
kilos. These, when roasted, are very tasty and nutritious. 
They are used in making confections and, by many, are con- 
sidered superior to almonds. 

The oil ^ from pili nuts has good keeping qualities, and a com- 
position similar to peanut oil. 

On account of the popularity of pili nuts the tree, Canarium 
ovatum, is called the pili-nut tree. The name "pili" is also 
applied locally to other Canarium species such as C. luzonicum. 
The latter also yields edible pili nuts but they are much smaller 
than those obtained from Canarium ovatum. 

Manila elemi is also known by the Spanish name, "brea blanca" 
(white pitch) . When the resin first exudes from the tree it is 
always soft and oily and has a very fragrant odor. On stand- 
ing, however, it gradually loses volatile constituents and har- 

Forest ranger Rabaya'' investigated the elemi industry in 
the province of Tayabas. Concerning Canarium luzonicum and 
elemi resin he states: 

Pili tree is deciduous. It changes its leaves once a year during the 
dry season. At the beginning of the dry season (in the month of Feb- 
ruary in the Province of Tayabas), some trees begin to shed their leaves. 
The falling of the leaves becomes general as the dry season advances, so 
that in the latter part of March and April most trees are leaflless or nearly 
so. Early in May, young leaves begin to grow and their purplish red color 
renders the trees conspicuous for miles. ...It appears to thrive best in 
Southeastern Luzon, where there are no distinct dry and wet seasons. It is 
also grown in the Islands of Masbate and Samar. In the Province of Ta- 
yabas alone, it is estimated that there are close to 25,000 pili trees avail- 
able for resin tapping. 

6 West, A. P. and S. Balce. Philip. Jour. Sci. 23 (1923) 29. 

7 Bureau of Forestry Report not published. 

"^8. 1 Manalo and West: Composition of Manila Elemi 113 

Tapping begins in the early part of July, or as soon as the new leaves 
become green and mature, and continues until March when the trees shed 
their leaves. The length of the tapping season is, therefore, a little over 
8 months. 

A young tree of 15 centimeters in diameter (B. H.) yields an average 
of 2 kilos of resin per month or 16 kilos per season, a 60 centimeter tree 
yields 60 kilos, while a tree 1 meter in diameter, 120 kilos or more. It 
is claimed that pili trees growing along the gulleys produce more resin. 

The greater bulk of the resin exported to foreign markets comes from 
Tayabas Province, principally from the towns of Gumaca, Calauag and 
Lopez. During the year 1925, a little less than 100,000 kilos of resin 
were collected in Tayabas Province alone, and if prices warrant exten- 
sive collection, it is believed that as much as 300,000 kilos can easily be 
produced in one season. 

In Masbate,^ according to Forester Zschokke, the trees are 
tapped at the beginning of the rainy season and the process is 
repeated every other day until December. The resin is col- 
lected once a month and one man can take care of 75 to 100 
trees. The trees must be visited regularly to get good returns. 
The resin can be gathered at almost all seasons, but towards 
the end of the dry season the flow is very slight. Collections 
are made from the same trees year after year. 

Clover says that he has seen large-sized trees on which at 
least ten pounds of the exudation had accumulated probably 
within a month. Bacon estimates that a mature tree will yield 
an average of 45 kilos per year. He says that he has seen 
as much as 32 kilos on a large tree. This amount repres- 
ented a two months flow. 

The volatile portion of the resin is called elemi oil and is 
usually obtained by steam-distillation. Clover collected sam- 
ples of elemi from a number of trees. These were distilled 
and the optical rotation, specific gravity, and refractive index 
of these distillates determined. The results varied consider- 
ably. Clover concluded that this variation was due to a dif- 
ference in the resin obtained from individual trees. He isol- 
ated various terpenes from his numerous samples. According 
to his investigations: 

The combined results obtained by a careful examination of the oils from 
21 individual samples of resin establish the true composition of elemi oil 
so far as these samples may be considered as representative of the ag- 
gregate product. In several cases, notably in the last sample examined, 
substances were obtained which were not encountered in any other; it 
seems possible, therefore, that were the investigation continued, still others 

8 West, A. P. and W. H. Brown. Philip. Bur. For. Bull. 20 (1920) 42. 

6961 8 

114 The Philippine Journal of Science ^^*^ 

would be found in which new constituents would appear, although such 
cases would be rare and the substances themselves would constitute so 
small a proportion of the aggregate oil that they would scarcely need to be 
taken into account. 

It is obvious that in considering Manila elemi or the oil obtained there- 
from as products of a species, we must deal with an aggregate sample of 
these products; a sample derived from so great a number of individual 
trees that the peculiarities of the individuals disappear. If the native 
gatherer of resin utilizes a large number of trees and regularly removes 
the resin from them in small portions, the product which he places upon 
the market will be nearly homogeneous and a representative sample; but 
if he obtains his resin from a limited number of individuals his product 
will not be representative and, if he utilizes resin which has accumulated 
upon the trees in large quantity, it will not be homogeneous. 

Granted that we have a representative sample of resin, the composition 
of the oil will also be influenced by the following factors: 

(1) The age of the resin. 

(2) The temperature of the distillation. This factor will largely de- 
termine the proportion of the high-boiling part of the oil and will influence 
the composition of the terpene portion, because some of the terpenes suffer 
a change at higher temperatures. 

(3) The length of time used in the distillation. This factor will in- 
fluence only the proportion of high-boiling oil. 

Yield of oil. — In the first seven samples examined considerable differ- 
ence was found in the oil content. While there may be a certain amount 
of variation shown by the individual samples in this respect, it is thought 
that the differences found are more directly connected with the age of the 
resin. As previously noted, Schimmel & Co. state that the yield of oil is 
from 15 to 30 per cent. In several cases where I have examined samples 
of fresh, soft, resin purchased in Manila, I have always found that total 
yield to be from 25 to 30 per cent of the weight of the resin. 

Bacon investigated the volatile oil separated from numerous 
specimens of elemi resin obtained from individual trees. His 
experiments verified the conclusions of Clover that elemi ter- 
pene oils vary a great deal in their boiling points and optical 

By the distillation of elemi in vacuo, Bacon obtained from 
12 to 18 per cent of terpenes and from 12 to 15 per cent of 
a higher-boiling oil. The distillation residue, usually amount- 
ing to about 70 per cent of the elemi, is a light-brown, trans- 
parent, solid resin, with a brilliant fracture. It is completely 
and easily soluble in the cold in the following solvents: al- 
cohol, ether, benzol, petroleum ether, ligroin, xylol, chloro- 
form, amyl acetate, acetone, methyl alcohol, carbon tetrachlo- 
ride, ehtyl acetate, turpentine, amyl alcohol, and glacial acetic 

^8» 1 Manalo and West: Composition of Manila Elemi II5 

acid. Bacon thought this product has commercial possibili- 
ties as a varnish gum. He subjected it to destructive dis- 
tillation. He says : 

One kilo of this residue was distilled in a 2-liter Jena flask. It melts 
quietly with very little darkening and no foaming, evolution of gas, or 
other signs of decomposition. As the temperature is increased, consider- 
able cracking and evolution of water soon takes place and the whole mass 
becomes very dark, the oil beginning to distill at a vapor temperature of 
200° C. The whole distillation is very quiet with no foaming or frothing 
and very little evolution of gas. The oil at first is green, then an intense, 
flourescent blue and toward the close of the distillation almost white. The 
total distillate was 670 cubic centimeters of which 40 were the aqueous 
layer. Three hundred and ten grams of black tar were left in the dis- 
tilling flask. The oil, shaken out with dilute alkalies, gave 18 grams of 
acids, while after saponification with alcoholic potash a further 7 gram 
of acids were obtained. The neutral oils distilled in vacuo gave fractions 
at 10 millimeters of about 90 grams each . . . 

There were no indications of any separable, individual products, and ni- 
tration and oxidation of these fractions under various conditions gave no 
promise of obtaining individual compounds. This oil obtained by the de- 
structive distillation of elemi is evidently a resin oil of the same general 
character and useful for the same purposes as other resin oils. 


The samples of Manila elemi used in this investigation were 
obtained from F. E. Zuellig Company (resin dealers), Ma- 
nila. The resin was collected in Tayabas province. 

Acid number. — The acid number was ascertained by treating 
approximately 1 gram of elemi resin with 50 cc. of a mixture 
consisting of neutral absolute alcohol and benzene (25 cc. each) 
and titrating directly with tenth-normal alcoholic potassium 
hydroxide using phenolphthalein as indicator. 

Saponification number, — For this determination approxim- 
ately 1 gram of elemi resin was treated with 50 cc. of absolute al- 
cohol after which 25 cc. of tenth-normal alcoholic potassium 
hydroxide were added. The mixture was heated on a stream 
bath with a reflux condenser for 2 hours. The solution was then 
cooled and the excess alkali titrated with tenth-normal sulphuric 
acid in the presence of phenolphthalein. 

Manila elemi analysis. — Preliminary experiments were car- 
ried out to ascertain a convenient method for separating the 
constituents of Manila elemi. The procedure outlined below 
gave satisfactory results. 


The Philippine Journal of Science 


Manila elemi (1) 

To elemi (25 g) add 800 cc. of alcohol. 
Warm on water bath. Allow mixture 

to stand overnight. Filter through 

a weighed filter paper. 


Dry at 
40'-50^ C, 
to constant 

Insoluble matter (2) 

Soluble elemi 
Volatile oil 
Nonvolatile oil 
Resenes, etc. 
Resin acids 

Steam distill. 

Volatile oil 

Extract with ether. 

Separate. Dry 

ether extract 

with anhydrous 

Na2S04. Filter. 

Distill off 

ether. Dry residue 


to constant weight. 

Residual mixture 
Nonvolatile oil 
Resenes, etc. 
Resin acids 

Volatile terpenes (3) 

Residual mixture 
Nonvolatile oil 
Resenes, etc. 
Resin acids 


Nonvolative oil 

Extract with ether. 
Separate. Dry ether 
extract with anhy- 
druous Na2S04. Filter. 
Distill off ether. 
Dry residue at 
40 °-50 " C, to 
4 constant weight. 

Nonvolatile terpenes (4) 

Resenes, etc. 
Resin acids 

Dissolve in ether. 
Shake with 1% aqueous 
KOH several times. 


Manalo and West: Composition of Manila Elemi 117 


Ether etxract 


Dry with 
Na2S04. Filter 
Distill off ether. 
Dry residue at 
constant weight. 

Aqeuous solution 
Potassium salts of 
resin acids 

Resenes, etc. (5) 

Steam distill to 
remove ether. 
Cool.9 Acidify 
with concentrated 
HCl. Filter. Wash, 
until free of 
chlorides. Drain 
and dry at 40 '-50° C, 
to constant weight. 

Resin acids (6) 

9 If a precipitate separates out at this stage proceed according to the 
following remarks on this outline of analysis. 

Insoluble matter. — Manila elemi (25 g) is treated with 300 cc. 
of alcohol (95 per cent). The mixture of elemi and alcohol is 
heated on a steam bath and shaken occasionally for about a 
half hour. It is then allowed to stand overnight to facilitate 
subsequent filtration. The insoluble matter consisting of for- 
eign material settles out on standing leaving a fairly clear su- 
pernatant liquid. The mixture is filtered through a weighed 
filter paper. The insoluble matter is washed with alcohol and 
the filtered washings are added to the soluble elemi solution. 
The filter paper with the insoluble matter is allowed to drain 
on layers of filter paper after which it is dried in an oven at 
from 40° to 50 °C., until the weight becomes constant. 

Soluble elemi, — The alcoholic solution of elemi is steam dis- 
tilled to remove the terpenes. This process usually requires 
about 20 hours to get most of the volatile terpenes. The small 
portion that steam distills with difficulty can be included in 
the nonvolatile terpenes that are subsequently obtained. 

The distillate (in portions of about 100 cc) is extracted with 
ether until it no longer has a milky appearance. Generally one 
extraction for each portion is sufficient. The combined ether 
extracts are dried with anhydrous sodium sulfate. The clear 
solution is filtered into a weighed flask and the ether distilled 
off. In order to eliminate the last traces of ether the flask 
is placed in an oven and heated for about 3 hours at a tem- 
perature of from 30° to 40°C., or until the weight becomes 
constant. The residue consists of the terpenes (3). 

118 The Philippine Journal of Science ^^^^ 

The residual mixture from the steam distillation is filtered to 
remove the solid residue. The filtrate, containing some non- 
volatile oil, is treated in the same manner as the volatile oil 
except that the last traces of ether are removed by heating at 
a temperature of from 40° to 50 ''C. The residue is the non- 
volatile terpenes (4). 

The solid residue is dissovled in ether and shaken several 
times with aqueous potassium hydroxide (1 per cent). The 
ether extract is separated from the aqueous solution and man- 
ipulated like the ethereal extract containing the nonvolatile 
terpenes (4). The final product contains resenes (5) and re- 
lated substances (amyrins, etc.). 

The aqueous solution from the ether extract is steam dis- 
tilled to remove ether and cooled to room temperature. A small 
amount of a white precipitate may sometimes appear at this 
stage as the separation of resenes and potassium salts of resin 
acids may not be complete. Ether is somewhat soluble in wa- 
ter and a small amount of resene constituents in the ether 
may remain with the aqueous solution of potassium salts. If 
a white precipitate forms it should be removed by filtering 
through a weighed filter paper. It is then dried to constant 
weight at from 40° to 50 °C. This weight is added to that of 
the resenes (5). 

By acidifying the aqueous solution of the potassium salts 
with concentrated hydrochloric acid the amorphous resin acids 
are precipitated. These are filtered through a weighed filter 
paper and washed with water until they are free of chlorides. 
The filter containing the acids is allowed to drain thoroughly 
after which it is dried in an oven at from 40° to 50° C, until 
the weight of the resin acids (6) becomes constant. 

Water in Manila elemd. — This resin usually contains a small 
amount of water, less than 5 per cent. The presence of water 
can be proved by adding to an interior portion of the resin 
some powdered anhydrous copper sulfate which immediately 
turns blue. Water is conveniently determined by difference, 
that is, the difference between 100 per cent and the sum of the 
percentages of all the other constituents contained in the elemi. 

Elemi constituents, — The data (Table 1) show that high- 
grade Manila elemi, containing only a small amount of insol- 
uble matter, consists principally of resenes together with some 
resin acids and volatile terpenes. It also contains a very small 
amount of nonvolatile terpenes. The resenes are not acted 
upon by alkalies and other common chemical reagents. This 

^8» 1 Manalo and West: Composition of Manila Elemi 119 

characteristic is said to enhance the value of resene resins 
for the manufacture of varnishes. 

Table I. — Composition 

of Manila elemi. 






Insoluble matter 

Per cent 






Per cent 






Per cent 


Volatile--- - 



Resenes, etc. 


Resin acids - _ - -_- - - 


lyfoisture (bv difference) - - - - --_ - «-- - - 


Total - . _ - — - 




All three samples had the same general constituents but 
the percentage of any particular one varied somewhat in the 
different samples. Specimens of elemi from the same tree may 
not have the same composition due to ageing and v^eathering. 
Resin that has been clinging to a pili (elemi) tree for some 
time and has become rather hard and weathered will naturally 
have a composition somewhat different from soft resin very 
recently exuded from the tree. 

The important constants of one of these samples (Table 1) 
are given below: 

Sample No. 2: 

Saponification No 28.53 

Acid No 18.50 

Ester No 10.03 

Here again these data show that Manila elemi contains only 
a comparatively small amount of free resin acids and saponi- 
fiable matter. 

M. Mladenovic^o analyzed a sample of Manila elemi by a 
method quite different from the one we used and obtained 24 
per cent of volatile oil, about 15 per cent of resin acids and 
about 60 per cent of resenes and related substances. These 
results are similar to the data given in Table 1. 

In accordance with the procedure in the text, the resin acids 
(6), were prepared from elemi resin (sample No. 3, Table 1). 
These acids gave the following constants: 

Saponification 125.00 

Acid No 108.47 

Ester No ^^'^^ 

10 Bull. Soc. Chim. Koy. Yougoslav. 8 (1937) 175. 

120 The Philippine Journal of Science ^^^ 

As shown by the data this product consists of a mixture of 
free acids and a saponifiable substance. 

During recent years considerable research has been done on 
the resene and resin acid portions of Manila elemi. Each of 
these was found to consist of a mixture of substances. Fur- 
ther investigations along these lines should be carried out. 


Manila elemi is a resin that is contained in the bark of the 
pili tree, Canarium luzonicum (Blume) A. Gray. This tree 
grows abundantly in low and medium altitudes in the primary 
forests of Luzon, Mindoro, Ticao and Masbate. 

When the bark of the pili tree is cut the resin oozes out. 
At first it is always soft and has a very fragrant odor. On 
standing, however, it gradually loses volatile constituents and 
hardens. The resin is said to flow best during the rainy season 
and accumulates in large masses of more than 30 kilos. It is 
used in making plasters, ointments, paints, varnishes, and for 
various other purposes. 

An accurate and convenient method was devised foif 
analyzing Manila elemi. This resin was found to consist 
principally of rescues together with some resin acids and 
volatile terpenes. By steam distilling the resin the volatile 
terpene oil passes over into the distillate leaving a residue of 
rescues and other elemi constituents. These various groups 
of residual substances are separated from each other by ap- 
propriate procedures. 

The rescues are not acted upon by alkalies and other common 
chemical reagents. This characteristic is said to enhance the 
value of resene resins for the manufacture of varnishes. 

Using a different method of analysis M. Mladenovic obtained 
results similar to those recorded in this report. 

78' 1 MaTialo and West: Composition of Manila Elemi 121 


Books reviewed here were received by Tlie Philippine Journal 
of Science from their respective publishers. 


Surface Active Reagent; Theoretical Aspects and Applications. By C. B. 
Young and K. W. Coons. Brooklyn, Chemical Publishing Co., Inc! 
1945. 381 pp. illus. Price, f6.00. 

Information obtained from this book will prove of great 
value to chemists and other technical men of industries. This 
book contains numerous tabulated data and illustrations, and 
a list of selected references at the end of each chapter. 

The first part of the book deals in detail with the theoretical 
side of surface tensions, the relation between surface tension 
and other chemical and physical properties of matter and 
the structures and effects of surface active agents. Methods 
of determining surface tension are given. The apparatus 
used are described. Chemical and mathematical equations 
and a comprehensive table of such data as the trade names of 
the wetting agents, type or chemical compositions of the com- 
pounds and their use in industry and manufacture are also 

The second part discusses extensively the utilization of the 
phenomenon of surface tension in various branches of in- 
dustries. Many specific formulas for emulsions and other 
products based on the applications of surface tension agents 
are given at the end of each chapter. — C. N. C. 

Fundamental Electronics and Vacuum Tubes.— By Arthur Lemuel Albert. 
New York, The Macmillan Co., 1947. 510 pp., illus. Price, }6.00. 

This book is devoted to a study of the fundamental principles 
governing the operation of electronic vacuum tubes and asso- 
ciated devices with the aim in view of imparting to the tech- 
nical student a thorough knowledge of the great possibilities 
which the vacuum tube may be put into use in the ever expand- 
ing field of electronics. 

The first chapter deals with the elementary concepts of 
matter, waves, electron, proton, positron, neutron, atomic 
structure, photon, quantum-treated in accordance with the 
latest theories and in a precise and clear style. This is 
followed in the next two chapters by the various processes of 
electron emission, the third chapter being devoted 'entirely to 
thermionic emission which is the most common type of electron 
emission used in commercial electronic today. The succeeding 

122 2^^^ Philippine Journal of Science *^' 

five chapters treat of the electrical characteristic of practically 
all types of vacuum tubes, from diodes to multi, electrode va- 
cuum, both gaseous and high vacuum types. The treatment 
is mathematical and includes a considerable number of cha- 
racteristic curves and diagrams which may be of practical 
value to the technical researcher. The last seven chapters 
are devoted to the applications of electronic vacuum tubes 
apparently with especial reference to communication pur- 
poses — ^the use of vacuum tubes as voltage amplifiers, oscilla- 
tors, including ultra high frequency oscillators, modulators, de- 
modulators, etc. A discussion on the use of vacuum tubes as 
measuring devices, such as vacuum tube voltmeters, cathode 
ray tubes and photo-electric tubes, is also included. 

References, questions and problems are provided at the end 
of each chapter, making the entire treatise a suitable textbook 
for any one who desires to have a thorough foundation in elec- 
tronics. — E. P. R. 

Dynamic Aspects of Biochemistry. By Ernest Baldwin. New York, 
The Macmillan Co., 1917. 456 pp., illus. Price, $4.00. 

This book gives a good idea about enzymes, with their 
functions and properties well discussed. The role they play 
in the utilization of food substances by the animal and human 
body is well presented. The book gives a good discussion on 
the ability of simple organisms to utilize simple substances 
for their needs, deriving their energy from the energy locked 
in complex substances. It also shows the dependence of 
higher forms of animal life for their subsistence on higher 
forms of animal and plant. 

Students on the fundamentals of fermentation should take 
note of this book for its well written accounts on carbohy- 
drates, nitrogen, proteins, fats and the end products of meta- 
bolism. This book is an excellent text or reference for students 
in fermentation. — R. R. 0. 

Inorganic Process Industries. By Kenneth A. Kobe. New York, The 
Macmillan Co., 1948. 371 pp., illus. Price, $6.00. 

The book is an able presentation of the processes involved 
in the operation of some inorganic chemical industries. The 
first chapter is devoted to technical literature on related sub- 
jects. It gives a practical approach to the first problem con- 
fronting a researcher or a technologist seeking information 
on matters pertaining to his field of activities. The other 
chapters give a thorough description of the manufacturing 
processes, the function of each equipment as a unit as well 

78. 1 Manalo and West: Composition of Manila Elemi 123 

as an integral part of the whole system and diagramatic 
illustrations ef the steps involved in the processing of mate- 
rials from their raw conditions to their final forms. From 
the accumulated information, so methodically presented and 
comprehensibly discussed, the researcher or professional man 
is placed in a position to understand thoroughly the conditions 
under which the different units of a system operate to attain 
the desired results at a minimum expense. 

The theoretical aspects of each process is also ably discussed, 
with a classified list of the references wherefrom detailed 
technological information on the various phases of the manu- 
facturing operations may be secured. 

The economic side has not been neglected. The relative 
costs of production by different processes and the marketing 
problems relative to trade names, commercial gradings, method 
of shipping, etc. of products are thoroughly discussed in the 
most practical way. 

The book as a whole is a valuable guide for technical men 
engaged in teaching, research, factory control or consulting 
activities and for students of industrial technology. It should 
form an integral part of their reference libraries. — R. H. A. 

Zinsser's Textbook of Bacteriology. Revised by David T. Smith and others. 
9th ed. New York, Appleton-Century-Crofts, Inc., 1948. 992 pp., 
illus. Price, $10.00. 

Zinsser's Textbook of Bacteriology is already known and 
has been used by medical students since its first publication 
in 1910. The 9th edition, 1948, of the book which was revised 
and edited by the students of the author gives in detail the 
modern conception of the biological characteristic of the organ- 
isms and the reactions of the living tissues to the bacteria and 
their products. 

The book is divided into seventy-seven chapters. The out- 
line and arrangement of the chapters remain the same as in 
the eighth edition with the exception of the chapter dealing with 
Pleuro-pneumoniae-like organisms and the Testing in vitro of 
sulfonamides and antibiotics (penicillin and streptomycin) 
sensitivity which are new additions. The nomenclature is based 
on Bergey's Manual of Determinative Bacteriology, 6th edition, 
1948. (For example in chapter XX'XIII Salmonella typhosa 
and Typhoid fever, the causative bacillus of the typhoid fever, 
had many names : Bacillus typhosus, Bacillus typhi abdominalis, 
Salmonella typhi, Eherthelta typhosa and finally Salmonella 

124 The Philippine Journal of Science 

Quite a long list of antibiotics is presented with their agent, 
source, range of activity, toxicity, properties and uses. 

The definition of every subject is simple and sufficiently 
comprehensive, and every discussion is clear and accurate 
based on up to date findings of known researchers. The book 
contains every phase of bacteriology, from bacteria, moulds 
fungi, filterable viruses, their environment, metabolites, modes 
of infection and control to immunology, antibodies and com- 
pliment fixation. In the discussion of the normal blood typing 
it also shows the importance of the detection of the Rh 
factor, which is important because of its immunizing ability. 

Extensive work on filterable viruses has been recorded and 
supported with electron micrographs. The medical mycology 
is fairly discussed with many recent illustrations of the cau- 
sative organisms. Due to the clear and accurate presentation 
of facts and the inclusive and thorough coverage of every- 
thing regarding bacteria known to date, the Zinsser's Text- 
book of Bacteriology is highly recommended for medical 
students, medical practitioners and health officers not only for 
their study but also as a guide for extensive work. The book 
is also valuable for the extensive references cited in it. — M. B. 

A Textbook of Heat for Upperclassmen. By LeRoy D. Weld. New York, 
The Macmillan Co., 1948. 436 pp., illus. Price, $5.00. 

A glance on the table of contents will show that the subject 
matter treated is adequately covered. The outstanding feature 
of the book is its plain and simple treatment of well-known 
theorems and laws; this style is well shown in Chapter I when 
the author made use of Longfellow's character in "Song of 
Hiawatha'' Pak-Puk-Keewis "who can pass readily from one 
another of a great variety of forms" to avoid confusion in the 
often use of expressions as transfer of heat, heat^ of vaporization, 
etc. And though he intended this textbooks for upperclassmen 
familiar with differential and integral calculus, he proceeded 
to show step by step the derivation of every formula — thus 
keeping the treatment well within the comprehension of the 
usual run of students. For those who want to delte deeper in 
the subject, a list of references is appended to each chapter. 
In short, here is your "Textbook of Heat" in eleven easy chap- 
ters. — L. G. A. 

The Philippine 
Journal of Science 





By RiiFiNO B. Gapuz and Domingo Santiago 

Of the Malaria Control Division, t/. S. Public 

Health Service, Manila 


King (1932), in his descriptive notes on the three PhiHppine 
Anopheles of the f unestus-minimus subgroup, described the larvae 
of Anopheles minimus flavirostris (Ludlow) and Anopheles 
mangy anus (Banks) as follows: 

Anopheles minimus var, flavirostris, — Clypeal hairs simple and unfrayed, 
outer and posterior occassionally forked; the posterior hairs as a rule do 
not extend beyond the base of the inner (with the head in a horizontal 
position) ; inner submedian prothoracic hairs usually with more than 21 
branches; leaflets of thoracic palmate tapered to a short point, without a 
filament; tergal plate of segment 1 narrow, oblong; plate of second segment 
deeply concave with the small median plate or a portion of the main 
plate detached ; tergal plates on segments 3 to 7 usually convex posteriorly 
and narrowed toward ends; anterior tergal hairs (hair 0) usually arise 
well away from the plate; antepalmate hairs (hair 2) of segments 2, 3, 
and 7 branched from the basal third; dorsal surface of larva with heavy 
subdermal pigmentation. 

Anopheles mangyanus, — Clypeal hairs similar to the above except that 
the outer and posterior are longer, the latter extending to or nearly to 
the edge of the clypeus; inner anterior prothoracic hairs usually with 
from 18 to 23 branches; leaflets of thoracic palmate hairs with a fine 

^This study was made under the auspices of the U. S. Public Health 
Service in connection with the Philippine Rehabilitation Act of 1946 of 
the 79th Congress of the United States. 

8104 127 

128 The Philippine Journal of Science ^^^ 

filament; tergal plates of segments 1 and 2 similar to var. flavirostris, 
those of segments 4 to 7 with broader ends; small submedian plates 
usually chitinized; anterior tergal hairs usually arise from the tergal 
plate, sometimes near the edge either just on or just off the plate; ante- 
palmate hairs of segments 2 and ^^ usually branched from the middle 
third; antepalmate hair of segment 7 simple, or occassionally forked 
apically, larvae pigmented with abundant spots of opaque white that are 
collected into submedian streaks on the thorax. 

Keys to hirvm of A. minimus fiavirostris and A. mangyanu^. — 
In 1934 Russell and Baisas made a key to the Philippine species 
of the genus Anopheles. The two species in question may be 
identified, thus: 

1. Inner clypeal hairs widely separated 11 

11. Inner and outer clypeal hairs simple 12 

12. Palmate 1 developed ^ 13 

13. Tergal plates large 14 

14. Leaflets of thoracic palmate hairs extended into long 

slender filaments ; antepalmate VII simple or forked 

apically mangyanus 

Leaflets of thoracic palmate hair not extended into 
long slender filaments; antepalmate ^11 branched 
basally minimus var. fiavirostris 

Russell, Rozeboom, and Stone (1934) adopted a key based 
on that by Russell and Baisas (1934), to wit: 

1. Inner clypeal hairs widely separated 12 

12. Clypeal hairs simple 13 

13. Abdominal segment 1 with developed palmate hairs 14 

14. Abdominal tergal plates large 15 

15. Thoracic palmate leaflets produced into long slender 

filaments; abdominal segment 7 with antepalmate 

hairs (No. 2) single or forked apically mangyanus 

Thoracic palmate leaflets not produced into long slen- 
der filaments; abdominal segment 7 with antepal- 
mate hairs (No. 2) branched basally minimus fiavirostris 

In 1946, Baisas published a key to the larvae of Philippine Ano- 
pheles based on natural groupings. It may be mentioned that 
this is now generally used in the identification of Anopheles 
larvae in the Philippines. 

,^ The three species under Group Myzomyia (inner anterior 
clypeal hairs widely separated, tergal plates large) may be 
differentiated as follows: 

1. Inner clypeal hairs with fine lateral branches; tergal plate II 

convex at the posterior border filipinse 

Inner clypeal hairs simple; tergal plate II concave at pos- 
terior border 2 

Gapuz and Santiago: Larvm of Anopheles 129 

2. Antepalmate VII simple or branched apically; leaflets of thoracic 

palmate extended into long points mangyanus 

Antepalmate VII branched near the base; leaflets of thoracic pal- 
mate taper into short blunt points minimus var. flavirostris 

Bohart (1945) in his A Synopsis of the Philippine Mosquitoes, 
describes the larvae of the above species as follows : 

Anopheles (myzomyia) mangyanus (Banks). — The larva is characterized 
by its simple clypeal hairs, the short and stoutly branched prothoracic 
hair 1, the slender but well developed palmate hair on abdominal tergite 
I and the rat-tailed apices of the leaflets on the palmate hair of tergite 
IV. It is very close to minimus flavirostris which differs in the somewhat 
thicker apices of the leaflets on the thoracic palmate hair and in having 
antepalmate hair 2 of abdominal tergite VII single or apically forked. 

Anopheles (myzomyia) minimus flavirostris (Ludlow). — The larva of 
this species closely resembles that of mangyanus but differs from it in 
having the apices of the thoracic palmate hair not drawn out into filaments 
and in having the antepalmate hair 2 of abdominal tergite VII branched 

It will be noted that of the three recognized Philippine species 
of the Group Myzomyia, Anopheles filipinse Manalang is easily 
identified by the convex posterior border of the second tergal 
plate (large). But considerable difficulty has been encountered, 
especially by a novice, in the identification of A. mangyanits 
and A, minimv^ flavirostris (both with concave posterior border 
of tergal plate II). The thoracic palmate (Plate 1~A) and 
antepalmate VII (Plate 1~B) are not easy to locate, especially 
as in most cases the specimens are examined alive. Moreover, 
the character of the thoracic palmate leaflets can be appreciated 
best only when the high power objective (X 450) is used. This 
involves a considerable loss of time and may lead to erroneous 
identification especially when one has to examine hundreds of 
larvae. When the thoracic palmate and the antepalmate VII 
are used as landmarks, even an experienced worker may have 
to take some time to arrive at a correct identification. 

Oftentimes the aforementioned anatomical parts are detached 
or mutilated, in which case the examiner encounters considerable 
difficulty. The keys published by Russell and Baisas, Russell 
et al, and Baisas, mentioned previously, cannot afford any relief. 

When confronted by this predicament, few resourceful exam- 
iners may resort to King's description of tergal plates III to 
VII, which are "usually convex posteriorly" in A. minimiis flavi- 
rostris, or have "broader ends'' in A. mangyantis. Russell and 
Baisas (1934) also described tergal plates IV to VII as rounded 

130 The Philippine Journal of Science ^^^^ 

or blunt at the edges in A. mangy anus, tapering towards the 
edges in A. minimus flavirostris. However, these and other 
authors, for some reason or other, made no mention of the 
specific importance of the tergal plate or plates, since they 
invariably point to the thoracic palmate and antepalmate VII in 
their keys to larvae of Philippine anophelines. 


It has been observed that the seventh tergal plate, not to 
mention the other tergal plates, has a constant and definite 
shape, and that the shape of the tergal plate in A, mangy anus 
differs from that in A. minimus flavirostris. The posterior 
border of the seventh tergal plate in A. mangy anus is more or 
less a straight line (Plate 2~C), while in A minimus flavirostris, 
it is convex or curved posteriorly (Plate 2-B). 

This observation was subjected to an extensive and rigorous 
test, in the laboratory and in the field, in the hope of finding 
a variation or exception. Hundreds of specimens were examined 
and invariably the identification made according to the charac- 
ter of thoracic palmate leaflets and antepalmate VII corresponded 
with the identification made according to the character of the 
posterior border of the seventh tergal plate. 

Carrying the test further, fifty A. mangy anus larvse and also 
fifty A. minimus flavirostris larvse were collected at difl'erent 
places and at different times and were individually bred out. 
Without exception, the imago identification was the same as the 
corresponding larval identification. 


It will be seen that by the tergal plates alone, the three species 
under Group Myzomyia could be definitely and rapidly identified. 
The advantage of the seventh tergal plate over the thoracic 
palmate and antepalmate VII cannot be overemphasized. The 
first is easily located, cannot be detached, and the character 
of the posterior border can easily be appreciated even if live 
larvse are examined as early as the second instar. On the other 
hand the thoracic palmate and antepalmate VII are easily de- 
tached or mutilated, and not easily located, and their charac- 
ters are not easily evaluated especially when the specimen is 
examined alive. 

A comparative illustration of the landmarks used in the iden- 
tification of the larvse of A. minimus flavirostris and A. mang- 

Gapuz and Santiago: Larvse of Anopheles 131 

yanus, both those adopted in keys to larvse and those described 
in this papers, are shown in Plate 3 and Plate 4. 

The following key to the larvse under Group Myzomyia, that 
is, the larvae with large tergal plates, may be adopted especially 
where rapid identification is desired. 

1. Tergal plate II with convex posterior border filipinse 

Tergal plate II with concave posterior border 2 

2. Tergal plate VII with convex posterior border. 

minimus flavirostris 
Tergal plate VII with more or less straight posterior border. 

mang yanus 


1. Baisas, F. E. Key to larvae of Philippine Anopheles. Mimeographed 

for teaching purposes. 1946. Manila, Bureau of Health. 

2. BoHART, R. M. A synopsis of the Philippine Mosquitoes. U. S. Naval 

Medical Research Unit No. 2, NAVMED 580. Bureau of Medicine 
and Surgery, Dept. of the Navy. 1945. 87 pp. 

3. King, W. V. Three Philippine Anopheles of the funestus-minimus 

subgroup. Philip. Jour. Sci. 48 (1933) 485-523. 

4. Russell, P. F. and F. E. Baisas. A practical illustrated key to larvse 

of Philippine Anopheles. Philip. Jour. Sci. 55 (1934) 307-336. 

5. Russell, P. F., L. E. Rozeboom, and A. Stone. Keys to the Anophe- 

line mosquitoes of the world. Philadelphia, The American Ento- 
mological Society, 1943. 152 pp. 


Plate 1 

Fig. 1. Thoracic palmate of A, mangyanus and A. minimus flavirostris, 
compared. Character of leaflets can best be appreciated by 
using high power magnification (X 450). 
2. Antepalmate VII of A. mangyamus and of A. minimus flavirostris, 
compared. Its relation to other structures, indicated. 

Plate 2 

Fig. 1, Tergal plates of A. filipinse. Second tergal plate with convex pos- 
terior border. 

2. Tergal plates of A. minimus .flavirostris. Second tergal plate with 

concave posterior border. Seventh tergal plate with convex pos- 
terior border. 

3. Tergal plates of A, mangyanus. Second tergal plate with concave 

posterior border. Seventh tergal plate with more or less straight 
posterior border. 

Plate 3 

Fig. 1. Landmarks (second tergal plate, thoracic palmate, and antepalmate 
VII) used in the identification of A. minimus flavirostris ac- 
cording to keys of various authors. 
2. Landmarks (second and seventh tergal plates) used in the identifi- 
cation of A. minimus flavirostris as described in this paper. 

Plate 4 

Fig. 1. Landmarks (second tergal plate, thoracic palmate, and antepalmate 
VII) used in the identification of A. mangyanus according to 
keys of various authors. 
2. Landmarks (second and seventh tergal plates) used in the identifi- 
cation of A. mangyamus as described in this paper. 


Gapuz and Santiago: Larvje of Anopheles.] 

[Philip. Jour. Sci., 78, No. 2. 

I I 


\ -^ 

\/^ \ 

. mcrtfyanus 

A.mimmw fUvirosiris 

» mcn^tji 

2 A.whiimm flmrci^iris 


Gapuz and Santiago: Larvae of Anopheles.] 

[Philip. Jour. Sci., 78, No. 2. 


2 3 

A.mimmus fkmrmhis A 



Gapuz and Santiago: Larv^ of Anopheles.] 

[Philip. Jour. Sci., 78, No. 2. 


/ \ 


.-> ! 




1 2 

A. minimus flavtro^iris 


Gapuz and Santiago: Larv^ of Anopheles.] 

[Philip. Jour. Sci., 78, No. 2. 



By Deogracias J. Cabrera 

Of the Malaria Control Division, 

U. S, Public Health Service, Manila 

In the Philippines naganol ^ is extensively used in the treat- 
ment and prophylaxis of equine and bovine surra, a disease due 
to infection with Trypanosoma evansi (Steel, 1885). The dose 
of the drug ordinarily advocated and employed by most veteri- 
narians is 0.5 gm. per 100 pounds of body weight, but there are 
those who claim that a large knockout dose should be used in 
order to effect Ehrlich's therapia magna sterilizans. 

Before and during the early part of World War II, the writer 
conducted a series of in vitro and in vivo studies on the mode of 
action of naganol and other trypanocidal drugs. The object was 
to determine whether the drug has a direct trypanocidal effect 
and if it is more rapidly lethal to the trypanosomes in bigger 
doses. Several investigators, using various species of trypa- 
nosomes, have postulated that naganol has no direct trypanocidal 
effect, but according to the available literature the mode of ac- 
tion of the drug against T. evansi has not yet been studied. 


The strain of Trypanosoma evansi used in the in vitro studies 
was originally obtained from a naturally infected Philippine 
pony and was maintained in the laboratory in guinea pigs. 
The trypanosomes used in the in vivo experiments were from the 
blood of four horses, of which Horses A, B, and C were naturally 
infected animals which were brought to the laboratory for 
treatment. Horse C was a case which had relapsed from a 
previous treatment (drug used presumably naganol) instituted 
in one of the veterinary hospitals in Manila; Horses A and B 
were primary cases. Horse D was infected with trypanosomes 
from an experimental rat. ^ 

lAlso known as Bayer 205, germanin, antrypol, suramin, moranyl, 

Forneau 309. 



The Philippine Journal of Science 


Exposure of T. evansi to naganol in vitro: 

Experiment 1. — On October 15, 1940, two drops of horse blood 
teeming with trypanosomes (100,000,000 per c.c.) were added 
to each of three tubes containing 5 c.c. of a medium composed 
of equal parts of horse serum and Locke's solution with 0.2 per 
cent glucose and 0.1 mg. naganol per c.c. of the serum-Locke 
solution. The same amount of infected horse blood was added 
to each of two tubes containing 5 c.c. of the serum-Locke- 
glucose medium but with no naganol. The mixtures were left 
at room temperature for a period of three hours, after which 
they were centrif uged and the sediments washed two times with 
0.85 per cent sodium chloride solution. A drop of the sediment 
was examined under the microscope and if found positive for 
actively motile trypanosomes, one-half of the remaining sedi- 
ment resuspended in 1 c.c. of the supernate was inoculated into 
a white rat. Three white rats, Nos. NR-1, NR-2 and NR-3, 
were each inoculated in this manner with the sediments of the 
test tubes containing naganol and two other rats, Nos. OR-4 
and OR-5, were inoculated with the sediments of the control 
tubes which contained no naganol in the medium. 

The number of trypanosomes inoculated per rat, in both the 
naganol and the control rats, was approximately 3,000,000 per 
100 gram body weight, assuming that one-third of the original 
number was either lost or killed during the process of centri- 

Results. — The results are shown in Table 1. Rats NR~1, 
NR~2 and NR-3, which were inoculated with the naganol- 
exposed trypanosomes remained healthy, and no trypanosomes 
were seen in their peripheral blood during a six-month-period 
of observation. Control rats OR-4 and OR-5, on the other hand, 
showed the presence of trypanosomes in their blood on the 
fourth day from the time of inoculation and died on the seventh 
and eighth day, respectively. 

Table 1. — Showing the effect of naganol in vitro on the infectivity of 
Trypanosoma evansi 


per c, e 
0.1 mg. 
0.1 mg. 
0.1 mg. 


I*ength of 
in vdro 

3 hours 
3 hours 
3 hours 
3 hours 

3 hours 

before inoculation 

very active 
veiy active 
very active 
very active 

very active 




Period of 


of rats 

6 months 
6 months 

6 months 

7 days 

8 days 


Positive on 4th day; 

died on 7th day. 
Positive on 4th day; 

died on 8th day. 

Cabrera: Naganol as a Trypanocidal Agent 137 

Exposure of T, evansi to naganol in vivo: 

Experiment 2, — On August 13, 1942, Horse A, whose blood 
was teeming with trypanosomes, was given 0.6 gm. of naganol 
per 100 pounds of body weight intravenously. This dose, which 
is higher than the usual initial therapeutic dose recommended 
by veterinarians in the treatment of surra in the horse, was 
calculated to give a blood concentration of approximately 0.14 
mg. of the drug per c.c. The calculation was based on the 
probable blood volume of the animal, assuming that the total 
amount of blood in the horse is 9.7 per cent (Welcher, cited by 
Dukes, 1942) of the body weight and taking 1.06 as the spe- 
cific gravity of horse blood. After the injection, blood from the 
animal was examined microscopically for trypanosomes from 
time to time during a period of twenty-seven hours. 

Blood samples were withdrawn from the horse three hours 
and twenty-seven hours after the injection. One drop from each 
sample, which was positive for actively motile trypanosomes, 
was added to 5 c.c. of 1 per cent sodium citrate solution and 
1 c.c. of the mixture was injected subcutaneously into a white 
rat (HR-6 was given the three-hour sample and HR~7 the twen- 
ty-seven-hour sample). The amount of naganol carried over 
with the horse blood into each rat was calculated to give a rat 
blood level of about 013 micrograms per c.c. 

Results.— "The results are given in Table 2. Actively motile 
trypanosomes were observed in the blood of Horse A during 
a period of observation which lasted for twenty-seven hours. 

Rat HR~6 was observed for a period of forty-seven days; 
Rat HR-7 for one hundred seventeen days. No trypanosomes 
were seen in the blood of these animals during the respective 
periods of observation. 

Table 2,— Showing the effect of naganol in vivo on the infectivity of 
Trypanosoma evansi 



cone, in 
horse blood 

Period of 


in vivo 

somes before 


Period of 


of rats 


Horse A 
Horse A 
Horse B 
Horse B 
Horse G 

Horse C 

0.14 mg. 
0.14 mg. 
0.14 mg. 
0.14 mg. 
0.14 mg. 

0.14 mg. 

3,13 hours 
26.88 hours 
2.00 hours 
3.00 hours 
2.17 hours 

3.25 hours 

very active 
very active 
very active 
very active 
very active 

very active 



47 days 
117 days 
36 days 
36 days 
15 days 

22 days 

Negative , , 
Positive on 9th day; 

died on 15th day. 
Positive on 11th day; 

died on 22nd day. 

138 The Philippine Journal of Science ^^*® 

Experiment S, — On October 21, 1942, another naturally in- 
fected animal (Horse B) was treated with naganol to give a 
blood level of about 0.14 mg. of the drug per c.c. Blood samples 
were withdrawn two hours and again three hours after the in- 
jection of the drug. One drop from each of these blood samples 
was added to 5 c.c. citrate solution and 1 c.c. of the resultant 
mixture inoculated into a white rat (HR~8 was given the two- 
hour sample and HR-9 the three-hour sample). 

Results. — The results are shown in Table 2. Actively motile 
trypanosomes were demonstrated in every blood sample of 
Horse B examined at different intervals during a period of 
twenty-one hours. 

No trypanosomes were seen in the blood of Rats HR-8 and 
HR-9 during a period of observation of thirty-six days. 

Experiment 4. — This experiment is the same as Experiment 3, 
except that the animal used (Horse C) was a case which had 
relapsed from a previous treatment with naganol. 

Results. — The results are shown in Table 2. Actively motile 
trypanosomes were found in the blood of Horse C examined 
at different intervals during a period of twenty-one hours. 

On the ninth day from the time of inoculation, Rat HR-10 
was found positive for trypanosomes in the peripheral blood 
and died on the fifteenth day; Rat HR~11 showed the organisms 
on the eleventh day and succumbed on the twenty-second day. 
The long prepatent period and the protracted course of the 
infection in these animals are unusual and seem to indicate that 
either the virulence or the integrity of the parasites has been 
altered in some way by exposure to the drug. 

Experiment 5. — On August 14, 1941, experimentally infected 
surra Horse D was given by the intravenous drip method 
massive doses of naganol in two installments. The first install- 
ment, at a dose of 0.7 gm. per 100 pounds of body weight, was 
administered continuously during a period of two hours; the 
.second, after an interval of three hours at a dose rate of 1.3 
gm. per 100 pounds of body weight, was given continuously 
for another period lasting two hours. The sum total of the 
drug given was approximately 2 gm. per 100 pounds of body 
weight, or four times the usual therapeutic dose. Calculated 
from the live weight of the animal, this was equivalent to a 
naganol blood level of about 0.48 mg. per c.c. 

After the administration of the first dose and again after 
the second dose, blood was withdrawn and examined for trypa- 

Cabrera: Naganol as a Trypanocidal Agent 139 

nosomes. Blood samples were also examined after twenty-two 
and twenty-eight hours from the time the first dose was given. 
Results. — The results are given in Table 3. After the admin- 
istration of the first and second doses, the blood of this animal 
was found swarming with trypanosomes ; twenty-two hours 
after the administration of the first dose only a few but still 
actively motile trypanosomes were present ; in the twenty-eight- 
hour sample no trypanosomes were found. (The results of this 
experiment will be reported in greater detail in another paper.) 


The Philippine Journal of Science 










































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Cabrera: Naganol as a Trypanocidal Agent j^j^ 


The results of the in vitro study reported in Experiment 1 
show that naganol (Bayer 205) has a disarming action against 
Trypanosoma evansi, for although the exposed trypanosomes 
were actively motile when inoculated to rats they failed to cause 
infection. Several investigators (Nauck, 1925; Reiner and Ko- 
veskuty, 1927; Issekutz, 1933 and Hawking, 1939), working 
with different species of trypanosomes, have called attention 
to this disarming action of the drug. According to Reiner and 
Kovestuky (1927), Jancso and Jancso (1934) and Hawking 
(1939), this property of naganol has an opsonizing effect which 
renders the trypanosomes more sensitive to phagocytosis by the 
reticulo-endothelial system. Culbertson's (1939) observation on 
the more prompt action of germanin against T. equiperdum in 
old compared with young rats, which he believes to be related 
to a difference in the phagocytic capacities of the cells of rats 
of different age groups, and Hawking's (1940) success in cul- 
turing trypanosomes from the blood of a patient whose plasma- 
germanin level was 0.42 mg. per c.c, all seem to point to the 
conclusion that naganol has no direct trypanocidal activity. 

The results of the in vivo experiments also point to the same 
conclusion, for it will be noted that the trypanosomes persisted 
in the blood of all the treated animals for twenty or more 
hours after the administration of the drug irrespective of dose 
size used. If the drug had a direct trypanocidal activity, a 
more prompt knockout effect could be expected by the use of 
large doses of it. This was not borne out by the results of 
Experiment 5 in which trypanosomes were seen in the blood 
of Horse D twenty-two hours after it was given massive doses 
of the drug. 

Although the trypanosomes obtained from the experimental 
horses were not washed before inoculation into rats, the amounts 
of naganol carried over from the former to the latter are 
believed to be negligible. At most, the concentration of the 
drug in the inoculated rats could not have been more than 0.135 
micrograms per c.c. of blood. Against highly virulent strains 
of T. brucei, Vierthaler and Boselli (1939) have found 1.3 mg. 
per 100 c.c. of blood (13 micrograms per c.c.) to be the smallest 
protective blood level of germanin (naganol) in the blood of 
rabbits. Against a strain of T. rhodesiense, Hawking (1939) 
has noted 0.2 mg. per 100 gm. rat (about 22 micrograms or 

142 The Philippine Journal of Science iws 

gamma per c.c.) i as the minimum effective dose. These amounts 
are 95 and 160 times greater than the calculated blood level of 
naganol in our experimental rats. 

In rats inoculated with the blood of Horse C, the presence 
of a naganol-resistant strain of trypanosomes remains to be con- 
sidered. It is to be recalled that Hawking (1939), using T. 
rhodesiense, has found germanin-resistant trypanosomes not to 
lose their infectivity upon incubation in vitro with as much as 
1 mg, of naganol per c.c. of medium. With normal trypano- 
somes he found 0.1 mg. per c.c. to be sufficient to render the 
organisms noninfective to mice. 

It is possible that Horse C, before coming to hand, had al- 
ready been treated for surra with doses of naganol too small 
to he effective but enough to allow the trypanosomes to gradually 
develop tolerance against the drug, as has been observed by 
Van Hoof, Henrard and Peel (1938a) with T. gambiense. It 
is also possible that drug resistance may have been acquired 
after repeated exposures of successive generations of the strain 
to subcurative levels of naganol in the blood of one or more 
hosts of the same or different species. For these reasons, the 
potential danger of producing drug-tolerant strains of T. evansi 
through the unintelligent and indiscriminate use of naganol as 
a prophylactic and as a treatment for equine and bovine surra 
in the field should be borne in mind. 


The effect of naganol against Trypanosoma evansi in vitro 
and in vivo was studied. 

Virulent trypanosomes exposed in vitro for three hours to 
a naganol (germanin) concentration of 0.1 mg. per c.c. were 
rendered noninfective to white rats. 

Blood swarming with trypanosomes obtained from two horses 
naturally infected with surra two, three and twenty-seven hours 
after the intravenous administration of naganol failed to infect 
white rats. 

Blood obtained from a naturally infected horse, which had 
relapsed from a previous treatment (presumably with naganol), 
two and three hours after the intravenous injection of naganol,' 
was infective to white rats which died after unusually long 
prepatent and patent periods. It is believed that the infection 
in this horse was due to a strain which was naganol-resistant 

1 Calculation in terms of blood level is by the present writer. 

Cabrera: Naganol as a Trypanocidal Agent 143 

due to previous unsuccessful treatment with subcurative doses 
of the drug or as a result of repeated exposures of one or more 
generations of the strain to ineffective levels of the drug in 
the blood of one or more hosts of the same or different species. 
The potential danger of producing naganol-resistant surra or- 
ganisms through the indiscriminate use of the drug in the field 
is pointed out. 

In all of the naganol-treated horses, the trypanosomes were 
found actively motile up to twenty-seven hours after the ad- 
ministration of the drug. The organisms, however, were found 
to have markedly decreased in number at about the twentieth 
to the twenty-sixth hour. 

The apparent lack of positive correlation between dose size of 
the drug and the rate of disappearance of the parasites in the 
peripheral circulation lends support to the hypothesis that 
naganol is indirectly trypanocidal. 


The writer is much indebted to Dr. Zacarias de Jesus, 
former chief of the Parasitology and Protozoology Division, 
Bureau of Animal Industry and now Professor and Head, De- 
partment of Parasitology and Protozoology, College of Veteri- 
nary Science, University of the Philippines, without whose help 
and encouragement this study could not have been accomplished. 


1. CULBERTSON, J. T. Studies on age resistance against trypanosome 

infections. IV. The activity of germanin (Bayer 205) upon T. equi- 
perdum infections in rats of different age groups. Amer. Jour. Hyg. 
Sec. C. 29 (1939) 7377. 

2. Dukes, H. H. The Physiology of Domestic Animals, Ithaca and New 

York, Comstock Publishing Co., Inc. 1942. xiv 721 pp., 168 figs. 

3. Hawking, F. Contribution on the mode of action of germanin (Bayer 

205). Ann. Trop. Med. and Parasit. 33 (1939) 13-19. 

4. Hawking, F. Culture of T, gambiense in blood from normal and 

infected persons. Ann. Trop. Med. and Parasit. 34 (1940) 31-34. 

5. IsSEKUTZ, B. VON Beitrage zuer wiekung des germanins. II. Arch. 

Exper. Path. u. Pharmak. 173 (1933) 499. (Cited by Hawking, 

6. Jancso, N. von and H. von Jancso. Microbiologische grundlagen der 

chemotherapeutische wirkung. I. Wirkungs-mechanismus des germa- 
nins (Bayer 205) bei trypanosomen. Zentralbl. f. Bakt., Orig. 132 
(1934) 257. (Cited by Hawking, 1939.) 

7. MCLETCHIE, J. L. The control of sleeping sickness in Nigeria. Trans. 

Roy. Soc. Trop. Med. and Hyg. 41 (1948) 445-470. 

8104 2 

144 The Philippine Journal of Science *®*^ 

8. Nauck, E. Unterschunger uber die wirkung des trypanosomenheil* 

mittels *Bayer 205'. Arch. f. Schiffs- u. Trop.~ Hyg. 29 (1935) 1. 

9. Eeiner, L. and J. Koveskuty. Ueber den wirkungsmechanismus von 

Bayer 205. Deutch. Med. Woch. 53 (1927) 1988. 

10. Van Hoof, L., C. Henrard, and E. Peel. The stability of Bayer 

205 resistance in Trypanosoma gambiense* Trans. Key. Soc. Trop. 
Med. and Hyg. 32 (1938a) 197-208. 

11. Van Hoof, L., C. Henrard, and E. Peel. Ann. Soc. Beige Med. Tr6p, 

18 (1938b) 143 (Cited by Mcletchie, 1948). 

12. Vierthaler, R. W. and A. Boselli. Die bedeutung kleinster germa- 

ninmengen im kaninchenblut als schutz gegen eime infection mit Tryp- 
anosoma brueei (The significance of the smallest dose of germanin 
in rabbit blood as a protection against infections with T. brueei). 
Arch. f. Schiffs- u. Trop,- Hyg. 43 (1939) 149-160. 




By AuRELio O. Cruz 
Of the Institute of Science y Manila 

Singkamas oil is obtained from the seeds of an hairy her- 
baceous vine known botanically as Pachyrrhizus erosus (Linn.) 
Urb. Recently the composition of the oil was determined and 
the results showed that this oil consists principally of glycerides 
of oleic, linoHc and saturated acids and is very similar in 
composition to kapok, rice, cottonseed, and peanut oils. 

Brown and West ^ give the following description of Pachyr- 
rhiztis erosus: 

Pachyrrhizus erosus is a rather coarse, somewhat hairy, herbaceous 
vine. The leaves are compound with 3 leaflets which are up to 15 cm. in 
length and 20 cm. in width. The flowers are pale blue or blue and white, 
2 to 2.5 cm. long and borne in racemes which are up to 45 cm. in length. 
The pods are about 10 cm. long, 10-12 mm. wide, flat, hairy and contain 
from eight to ten seeds. The roots are large, fleshy, turnip-shaped. They 
are eaten raw or prepared in a variety of ways. The young fruits are 
sometimes eaten as a vegetable. 

This species is a native of tropical America, but is now widely dis- 
tributed in the tropics. It is thoroughly naturalized in the Philippines and 
is common in thickets. It is also extensively cultivated. 

Heyne ^ stated that Greshoff found in the seeds 38.4 per cent 
of a colorless, limpid oil. 

Lately several articles, dealing on the insecticidal property of 
Pachyrrhizus erosus beans, also known as yam beans, either 
alone or mixed with some other rotenone bearing plants were 
published. The results indicate the possibility of utilizing yam 
beans as insecticides. 

L. B. Norton » obtained by extracting the ground yam beans 
with ether, 1.4 per cent of the msecticidally active resin, after 
removing the oil. 

ip. I. Bureau of Forestry, Bull. 20 (1920). 

^De Nuttige Planten van Nederlandsch-Indie 2 (1916) 346. 

Bjour. Am. Chem. Soc. es (1943) 2257-60. 



The Philippine Journal of Science 


Shui-Luen Hwang ^ reported to have obtained 0.56 to 1.01 per 
cent of rotenone from twelve samples. He further stated that 
oil expressed from the seeds contained rotenone to an extent 
to make it of insecticidal value. 


The singkamas seeds used in this investigation were supplied 
through the kindness of Director Felix D. Maramba of the 
Bureau of Plant Industry. The seeds were ground to fine 
powder. The composition of singkamas seeds compared with 
soy beans is given in Table 1. 

Table 1. — Composition of Philippine singkamas seeds compared with 
Philippine soy beans. 




Soy beans 


Per cent 

9. a 








Oil (ether extract) 




Crude fiber 


Ash „ - 


Carbohydrates (by dif f .) _____ ___ . 





As shown by the data singkamas seeds have a high protein 
content like soy beans. 

Singkamas oil was prepared by extracting the ground seeds 
with ether. The extracted oil was purified by treating it with 
about 2 per cent of activated carbon. This treatment removes 
vegetable fibers and colloidal matter and produces a brilliantly 
clear yellowish brown oil. 

The constants of this sample of Philippine singkamas oil are 
given in Table 2. 

Table 2. — Physical and chemical constants of Philippine singkamas oil. 

Specific gravity at 31V4*'C 0.9186 

Refractive index at 31 "C 1.4730 

Saponification value 182.9 

Acid value 2.6 

Iodine number (Hanus) 84.1 

Thiocyanogen value 56.8 

Unsaponifiable matter 2.2 

The saturated and unsaturated glycerides in singkamas oil 
were calculated from the iodine and thiocyanogen values in 

4Kwangsi Agri. 2 (1941) 296-80 

Cruz: Philippine Singkamas Oil 


accordance with the Official and Tentative Methods of the Amer- 
ican Oil Chemists' Society. The results are recorded in Table 
3. Included in Table 3 are compositions of other Philippine 
and American oils for comparison. As shown by the data 
Philippine singkamas oil is very similar in composition to rice, 
kapok, cotton seed and peanut oils. All these oils consist prin- 
cipally of glycerides of oleic, linolic and saturated acids. 

Table 3. — Comparison of Philippine singkamas oil with other oils. 

Philippine oils. 

American oils. 


Kapok seed 

Rice oil 


Cotton seed 


Glycerides of: 

Unsaturated acids — 

Oleic - - - 

Per cent. 



Per cent. 



Per cent. 

Per cent. 

Per cent. 


Linolic. -- 

27.7 1 41.7 


Saturated acids — 
Unsaponifiable matter 







98.0 1 99.8 



a Philippine kapok-seed oil iCeiba pentandra Gaertner), analyzed by A. O. Cruz and A. 
P. West. Philip. Jour. Sci. 46 (1931) 131. 

t> Philippine rice oil (Hambas variety), analyzed by A. O., Cruz and A. P. West. 
Philip. Jour., Sci. 47 (1932) 487. 

c American cottonseed oil analyzed by G. S. Jamieson and W. F. Baughman. Jour. Am. 
Chem. Soc. 42 (1920) 1197. 

<» American peanut oil (Spanish type), analyzed by G. S. Jamieson, W. F. Baughman 

and D. Brauns Jour. Am. Chem. Soc. 43 (1921) 1372. 


Data recently published by several investigators showed the 
possibility of utilizing singkamas seeds as a source of insecti- 

Singkamas seeds have, like soy beans, a high protein content. 

In this investigation the composition of Philippine singkamas 
oil obtained from the seeds of this plant by ether extraction 
was determined. The results indicate that the oil consists prin- 
cipally of glycerides of oleic, linolic and saturated acids. Sing- 
kamas oil resembles kapok, rice, cottonseed and peanut oils, 
although in different proportions. 


By Albert W. C. T. Herre 
Of the School of Fisheries, University of Washington 

The writer completed his Checklist of Philippine Fishes in 
May, 1948. This work lists 2,145 species actually known within ' 
the political boundaries of the Republic of the Philippines. Were 
one to include the names of those fishes recorded from nearby 
localities within 10 to 100 miles of Philippine soil, and certain 
to be captured in Philippine waters later on, the number would 
reach 2,300 or more. Intensive collecting almost anywhere in 
the Philippines will bring additions to the fish fauna of the 

One of the least known regions of the Philippines, as far 
as its fish are concerned, is the east coast of Luzon, north of the 
latitude of Manila. I, therefore, take pleasure in presenting the 
description of a new labrid from that region. With it ai-e also 
some notes upon a few other fishes, two of them new to the 


Dorsal XII~8^ anal III-9; pectoral 11-13; scales in the lateral 
line 26, plus 3 on the caudal base; the tubules of the anterior 
half of the lateral line are branched, becoming simple posteriorly. 
Three scales above the lateral line to the first dorsal spine, 
9 below. Dorsal and anal each with a high basal scaly sheath. 
8 predorsal scales; a band of smaller scales behind the inter- 
orbital region above the posterior part of the eye, followed by 
a narrow naked area. 

The form is oval, the dorsal and ventral profiles both uni- 
formly convex, the depth 2.3 times in the length. The head 
is rather small, 2.8, the truncate caudal 3.5 times in the length. 
The large prominent eye is high up, 3.85 times in the head, 
its diameter a little less than the width of the slightly convex 
interorbital, 1.46 times in the snout, which is 2.63 in the 
head. The maxillary equals the interorbital, its posterior angle 
extending back below the anterior half of the eye, 3.57 in the 


150 The Philippine Journal of Science ^®*® 

head. The canines are prominent, the outer pair of the mandible 
much the largest; no posterior canine. The preopercle with 9 
rows of scales, its posterior and inferior limbs naked, its 
posterior border smooth; the opercle with 4 rows of scales, the 
two most posterior much the largest. The scales forming the 
posterior row at the caudal base enlarged and pointed. 

Dorsal and anal relatively low, their real height concealed by 
the high basal sheath, the five posterior spines equal to or slight- 
ly longer than the snout ; the membranes between the spines are 
deeply incised and prolonged in a filament extending beyond 
each spine. The soft dorsal equals the spinous in height. The 
first anal spine equals the eye, the third equals the snout, the 
second is nearly as long ; the anal rays like the soft dorsal. Pec- 
torals broad, the upper rays elongate, pointed, 1.16 in the head ; 
ventrals short, not reaching the anus, 1.78 in the head; least 
depth of caudal peduncle 2.27 in the head; upper caudal rays 
slightly produced. 

In life the general color dorsally was dusky, with a broad 
dusky or blackish band covering the posterior part of the body 
from the anterior part of the soft dorsal to the anal and back 
to the caudal base. Six brilliant orange bands with very 
dark blue margins girdled the body; the first descended from 
the eye, the second from the nape, the third from the dorsal 
origin, the sixth from the last dorsal spine and first dorsal 
ray. The lower part of the head and body was pale to whitish. 
Across the snout were two orange bands with indigo margins, 
and a broad orange band connected the eyes. The strong canine 
teeth were bright blue. The dorsal and anal each had a broad 
longitudinal orange band with deep indigo margins, their spines 
bright blue. 

The tips of the dorsal filaments and margin of the soft dorsal 
and anal were indigo. The pectorals were clear yellow, the 
ventral spines blue, the rays orange. The caudal fin had a 
broad basal yellow band, the rest of the fin colorless. 

In preservative, the orange cross bands blackened on the 
dorsal region but faded below, while their margins and the 
stripes and band on the head also blackened. 

Here described from the type and sole specimen, 139 mm. 
long, obtained on a coral reef at Baler, Quezon (Tayabas) 
Province, Luzon. This fish is notable for its brilliant colors 
and striking color pattern. It was collected by Mr. R. B. Fox, 
anthropologist with the Philippine National Museum. 

Herre: Philippine Fish Records 15 j 

Key to Philippine Species of Choerodon 
A. Ventrals greatly produced, extending upon anal in adults; silvery 
spot below middle of spinous dorsal, with a dark patch below it. 

C ohlig acanthus 
AA. Ventrals not reaching anal. 

B. Body more elevated, depth 2.3 to 2.6 in length. 

C. Six transverse orange bands with deep blue margins on 

head and body C. balerensis 

CC. No pattern of cross bands. 

D. Pale abdominal color extends upward as wedge or 

bar to lateral line below middle of spinous dorsal; 

caudal peduncle with pale saddle C. anchorago 

DD. An orange or yellow band from pectoral axil to 

caudal base C. zamboangx 

DDD. Pale line from snout through eye and small white 

spots on anterior scales of body. 

C. margaritiferus 
BB. Body slenderer, depth 2.5 to 3.3. 

E. A broad dark band on side of body. 
E. A dark band from axil to pectoral to posterior 
part of dorsal, widening posteriorly; a broad 

silvery band above it C. zosterophorus 

EE. A broad curved dark band from above pec- 
toral tip to posterior part of soft dorsal. 

C, melanostigma 

EE. No broad dark band on side; a dark spot 

below 3 posterior dorsal spines; a large pale 

blotch between soft dorsal and lateral line. 

C. schoenleini 



Catulus torazame Tanaka Jour. Coll. Sci. Tokyo 23 (1908) 26, Plate 
2, Fig. 2. Misaki, Sagami, Japan; Garman, Memoirs Mus. Com- 
parative Zoology 26 (1913) 77, Misaki, Japan. 

Halaelurus torazame Tanaka, Fishes Japan 5 (1912) 87; Schimdt, 
Rendu, Acad. Sci., U.S.S.R. (1930) 285, Figs. 1 and 2, Tokyo, 
Fusan, Misaki; Schmidt, Copeia, No. 2 (1930) 48, Figs. 1-3. 

Scylliorhinus torazame Fowler, U. S. Nat. Mus., Bull. 100, 13 (1941) 
36; Japan; Korea; after Tanaka. 

Scylliorhinus rudis Pietschman, Anzieg. Akad. Wiss. Wien 45 
(1908) 133, Japan. 

Halaelurus rudis Tanaka, Fishes Japan 1 (1911) 13 Fig. 12. 

This little catshark has hitherto been known only from Japan 
and Korea, and is now recorded for the first time from the 

152 The Philippine Journal of Science ^^' 

Among the sharks purchased at the Dumaguete market for use 
in the zoology class at Silliman University, Professor Guillermo 
Magdamo noticed some that were unlike any he had ever seen 
before. On examination I found them to be typical specimens 
of the white-spotted catshark of Japan, from 225 to 330 mm. 
in length. These sharks were taken off Dumaguete, Negros 
Oriental Province. 

This little shark may be recognized by its numerous small cir- 
cular white spots or dots, sprinkled on a gray background. 

This is an altogether unexpected addition to the Philippine 
fish fauna. As Professor Magdamo drily remarked, "It is a 
part of the Japanese invasion.'' 

ACHIROPmCHTHYS KAMPENI (Weber and de Beaufort) 

A specimen of this little known snake eel, family Ophich- 
thyidae, was obtained from the Mantugil River, 42 kilometers 
from the sea, in the Municipality of Villar, Zambales Province, 
Luzon. It is known to the Negritos there as "banota.'* 

This is the fourth specimen known, and the second Philippine 
specimen. It was previously recorded from the Pinacanauan 
River, at Barrio Lamug, Municipality of Peiiablanca, Cagayan 
Province, Luzon. This is over 100 kilometers from the sea. It 
was originally described from Humboldt Bay, New Guinea, near 
the mouth of the Mbai River. I also collected it in the Papenoo 
River, Tahiti, about a kilometer from the sea. 

The Zambales specimen is 395 mm. long, the depth 26.3 mm., 
the head almost 8 times in the length. The eye is about 20 
times in the head, and 2.5 times in the noticeably narrow and 
pointed snout. The lateral line is conspicuously marked by a 
row of circular yellow spots surrounding the pores. 

The specimen agrees very well with the descriptions of other 
specimens, but has 5 canines on the vomer instead of 3 or 4. 



Centropritis hirtmdaceus Cuvier and Valenciennes, Hist. Nat. Poiss. 
1 (1831) 450, Japan; Schlegal, Poiss., Fauna Japan (1842) 14, 
Plate 5, fig. 1, Nagasaki. 

Chelidoperm hirundacea Boulenger, Cat. Fishes, ed. 2, 1 (1895) 305, 
Japan and Arafura Sea, off the Kei Islands. Jordan and Richard- 
son, Proc. U. S. Nat. Mus. 37 (1910) 464, Suruga Bay and Sagami 
Bay, Japan; Tanaka, Fishes Japan 12 (1912) 343, Plate 92, figs. 
296 and 297, Nagasaki to Tokyo. 

Herre: Philippine Fish Record 153 

Centropristis pleurospilus Giinther, Reports Challenger Exped 
Zoology 1 part VI, Shore Fishes (1880) 37, Plate XVI, Fig. D 
Kei Islands. * * 

Three examples, 53, 86, and 105 mm. in length, were taken by 
trawl at the entrance to Manila Bay, in 65 fathoms of water. 
Dorsal X-10; anal III-5 or 6; scales in lateral line 44. Our spec- 
imens all have a row of 5 black spots along the side; in the 
smallest specimen they form an almost continuous bar. With 
age the spots separate and eventually disappear in adult life. 

This is another interesting addition to the Philippine fish 
fauna. Previously it has been known only from Japan and off 
the Kei Islands, which lie between New Guinea and Australia. 



This little severely stinging catfish is known from India 
through the East Indies to the Philippines. It seems to be rather 
common throughout its range, and is primarily a reef fish of 
shallow coastal waters. In the Philippines at least, it enters 
rivers and travels considerable distances from the sea. 

Long ago I obtained it from the Tanjay River, Negros Oriental 
Province, less than a kilometer from the sea. In 1940, in com- 
pany with Mr. P. R. Manacop of the Philippine Bureau of 
Fisheries, I collected it in a shallow rapid in the Cagayan River, 
Misamis Oriental Province, Mindanao, about 15 kilometers from 
the sea. In July, 1947, Mr. R. B. Fox, anthropologist of the 
Philippine National Museum, obtained 4 specimens^ from the 
Baiigan Labaw River at Villar, Zambales Province, Luzon. This 
locality is in the Zambales Mountains, 40 kilometers from the 
China Sea. These fish ranged in length from 59 to 89 mm. 
Specimens 85 to 89 mm. in length were several times as bulky 
as the shortest specimen. 

This stinging fish is well known to the Negritos of the Zam- 
bales Mountains, who call it *'napo''. As scorpion fishes are 
characteristically marine reef-dwellers, it seems noteworthy that 
this species should leave the sea to dwell in hill streams. Ap- 
parently they like to live in shallow rapids, where their dark 
mottled forms blend indistinguishably with the volcanic gravel 
and stones that strew the bottom of the stream. 


Weber and de Beaufort, Fishes Indo- Australian Archipelago 3 (1916) 

316, figs. 150 and 151. 
Herre, Philip. Jour. Sci. 24 (1924) 108. 
Herre, Field Mus. Natural History, Zoological Series 21 (1936) 38. 





By L. M. YuTUC and Herman Sher 

Of the College of Veterinary Science 

University of the Philippines 


Since trypanosomes are blood-inhabiting protozoan parasites, 
most writers tend to take the attitude that the flagellates 
in an infected animal are found in all the body juices. They 
are said to be present in enormous numbers in one part 
of the body while but a few in another. Animals having many 
parasites in the periph^al circulation, when killed show them 
also in the organs. However, if they are not demonstrable 
by microscopic examination in the former, they are also not 
found in the latter. The blood of animals suffering from the 
disease is always infectious by animal inoculation, although 
there are periods during its course when the trypanosomes 
can not be found for days by microscopic examination of the 
peripheral blood. Plimmer and Bradford (1889) and others 
working with tsetsefly trypanosomes state that the lympha- 
tics near the point of inoculation first show the parasite and 
that the blood of the animal may be infectious for two days be- 
fore the parasites are found in it. 

According to Voges (1901) the whole cycle of the para- 
site of "mal de caderas," a species of trypanosome of the evansi 
group, takes place in the blood; and it would seem that the 
experiments conducted to determine the place of multiplica- 
tion in the body of the host tend to substantiate this state- 
ment. It is true that the trypanosomes are pretty evenly dis- 
tributed in the body juices and that similar forms are found in 
all parts of the body. Further, Musgrave and Clegg (1903) 
state that when an animal, in the blood of which trypanosomes 
are present, dies, the organisms are then found in all organs; 
and conversely, where none appear in the former, they are 
also absent in the latter. They are generally distributed and 
multiplication forms do not appear in exceptional numbers in 


156 Tlie Philippine Journal of Science *^^^ 

any organ. However, they are usually somewhat more num- 
erous in the spleen, liver and the lymphatic glands than in the 
bone marrow, and are seldom present in the medullary canal. 
They are found in the serous fluids and exudate of the joints, 
but rarely "in the urine". More recently, investigation in cat- 
tle and water buffaloes conducted by Manresa and Gonzalez 
(1936) showed that smears taken from all important organs of 
the animals with heavy peripheral infection at the time of kill- 
ing, invariably showed the presence of surra organisms in the 
heart, liver, spleen and lungs. In other cases, however, the 
results were variable. 

On the other hand, there are some investigators who differ 
with the general statement given above. They believe the 
parasites to be more numerous in certain organs, such as the 
lymphatic glands and the bone marrow, than in others. The 
results of the investigations of Schelling (1901) disclose that 
Trypanosoma evansi was not found in the spleen when it was 
present in the peripheral blood, and that the trypanosomes 
might be absent from the fluids and tissues but was constantly 
present in the bone marrow. The peritoneal exudate and bone 
marrow showed the parasites in the "budding form". The num- 
ber of trypanosomes in the spleen varied greatly. The conclu- 
sion was drawn that the parasite undergoes multiplication in 
certain organs, while it is destroyed in others. Martini (1903) 
regards the spleen, lymphatics, bone marrow, and, to a less ex- 
tent, the liver and the kidneys as the sites for the destruction of 

Citing the findings of Yorke (1911), Wolback and Binger 
(1912), and Stephenson (1922), Craig and Faust (1937) state 
that in African sleeping sickness, T. gambiense is found in the 
blood and lymphatic glands during the early or febrile stage 
of the disease. After the development of nervous symptoms 
the organism is found in the cerebrospinal fluid. It does not 
live within the tissue cells but may be found in the connective 
tissue spaces of the various organs and in the reticular tissue of 
the lymph nodes and spleen. It may be found in the intercel- 
lular spaces in the brain and is present in large numbers in 
the lymph channels throughout the body. Yutuc (1934) has 
demonstrated the presence of T. evansi after its prepatent pe- 
riod in the cerebrospinal fluid of horses by subinoculation and 
microscopic examination as early as five days and as late as 
twenty two days with an average of twelve days in ten out of 
seventeen animals. 

Yutuc and Sher: Distribution of Trypanosoma evansi 157 

In view of the divergent findings of the investigators cited 
in the foregoing, the present work was conducted with the fol- 
lowing objects: First, to determine the distribution of surra 
trypanosomes in the body of rats during the prepatent period; 
Second, to estimate the degree of trypanosome density in var- 
ious organs as well as in the blood during the patent period; 
Third, to determine whether or not the trypanosomes tend 
to appear in considerable numbers in some internal organs dur- 
ing periods when they can not be seen by the microscope in the 
peripheral blood (subpatent period) of guinea pigs. 


Animals used. — ^White and piebald rats and guinea pigs were 
employed in the investigation. The rats and a number of 
guinea pigs were obtained from the rat and guinea pig 
colonies of the Department of Bacteriology and Pathology, 
College of Veterinary Science, University of the Philippines, 
Pandacan, Manila. The rest of the guinea pigs came 
from the stock kept in the Department of Veterinary Medicine 
which have been raised from a single pair secured from 
the Institute of Hygiene, University of the Philippines. 

Animal care. — The rats and guinea pigs were kept in gal- 
vanized iron cages with movable pans. All pans were cleaned 
once a week. The animals were supplied with abundant feed 
consisting of copra meal and first class rice bran. Green for- 
age, mostly guinea grass was given in unlimited amount. 
Clean tap water for drinking was accessible all the time. 

Strain of trypanosome used.—lihe organism, Trypanosoma 
evansi (Steel, 1885) used was recovered July 23, 1946 by sub- 
inoculation into a guinea pig from a water buffalo suffering from 
surra kept for sometime in the animal shed of the Bureau 
of Animal Industry, Pandacan. The organism has been main- 
tained in the laboratory by continued passing through guinea 
pigs at appropriate interval. 

Procedure employed.— hi the first series of rats used in the 
study of the pre-patent period, infected blood was taken from 
the marginal ear vein of a guinea pig suffering from surra. 
The blood was aspirated in a one-cubic-centimeter syringe 
containing physiological saline solution to which a small amount 
of sodium citrate was added to prevent coagulation. One 
fifth cubic centimeter was inoculated subcutaneously into each 
rat. Inoculations of subsequent series of rats were made from 

158 The Philippine Journal of Science *^*^ 

the heart blood of the infected rats killed in the course of the 
study. Before the parasite appeared in the peripheral blood, 
the rats were killed with chloroformi. On autopsy impression 
smears of the important solid organs of the body were made; 
viz. — bone marrow, heart, liver, lungs, brain, kidneys, spinal 
cord and precural lymph glands. Occasionally other organs 
were included as may be noted elsewhere in this paper. In 
the process of posting the animals, care was taken in using 
clean instruments to avoid contamination of the smear of one 
organ with that of another. The average time consumed in 
the preparation of smears was approximately one half hour. 

The smears were dried and fixed in air. The preparations 
were then immediately stained with Wright's stain and dried. 
Each slide was examined microscopically for surra organism. 
At least fifty microscopic fields were examined and the average 
per field computed. 

In the study of the patent period the course of the disease 
was allowed to progress until the trypanosomes could be dem- 
onstrated by microscopic examination of the tail blood. In 
most instances the rats were bled to death or chloroformed 
when the blood was teeming with the causative agent. In 
the case of the subpatent period, the guinea pigs were the ani- 
mals of choice, because of their resistance to surra trypano- 
some. The course of the infection is of a relapsing type, the 
guinea pig dying as a rule after a number of relapses, hence 
appropriate for this study. In the preparation of the smears, 
the same routine was observed as the prepatent period of 

In the evaluation of the findings, the following key was 
employed : 

= No trypanosome in 50 microscopic fields 

+ = Less than one trypanosome per microscopic field 
2-f = From one to five trypanosome per mocroscopic field 
3-|- = From 5 to 80 trypanosomes per microscopic field 
4+ = Over 80 trypanosomes per microscopic field 


Prepatent period. — Table 1 shows that among the organs 
consistently examined for trypanosomes only the bone mar- 
row failed to yield the organism, while the brain, heart, kid- 
neys, and spleen showed the parasite once, the liver twice and 
the lungs three times. 

YutUG and Sher: Distribution of Trypanosoma evansi 159 

In the case of the other organs, which were sporadically 
examined, the organism was never demonstrated microscopi- 
cally. The data presented indicate that during the prepatent 
period the organism is also scarcely seen in the various solid 
organs examined. This agrees with the finding of Voges 
(1901) that the whole cycle of T. equinum, the causative agent 
of "mal de caderas," takes place in the blood of the vertebrate 
host. The occasional finding of the trypanosome in the or- 
gans examined can not be interpreted to mean that the 
parasites multiply and accumulate in considerable number in 
the internal organs during the initial period of the infection. 
Further, the morphological studies of the few organisms en- 
countered did not show the presence of dividing forms. 

Table 1. — Distribution of trypanosomes in the body of rats during 
pre-patent period of the infection 

Organs examined 

Rat number 



























Bone marrow -_ 

Heart --- 



Liver - 

Lunes ------ 

Spleen - 














s kil 

ed a 




* Mouse 

Patent period. — In the first series of rats as shown in Table 
2, the animals were bled to death at the height of the infection, 
that is when the peripheral blood was teeming with trypano- 
somes. Since it was impossible to examine all the animals at 
the same time, some of the killed rats had to be kept in the 
refrigerator for as long as two days. The distribution of the 
surra organisms appears decidedly different in the various or- 
gans examined. The heart and liver consistently showed heavy 
trypanosome burdens ranging from 2+ to 4+ in the former 
and 3+ to 4+ in the latter. The kidneys and the lungs come 
next with a trypanosome density of + to 4+. In general, the 
brain, bone marrow, and the spleen showed light trypanosome 
contents (0 to +). It is apparent from these findings that the 

8104 3 


The Philippine Journal of Science 


more vascular the organ the heavier is the trypanosomes density. 
The observation of Musgrave and Clegg (1903) that the surra 
organisms are usually somewhat more numerous in the spleen, 
liver and lymphatic glands than in the bone marrow is therefore 
not entirely confirmed. 

Table 2. — Distribution of trypanosomes in the body of rats at the height 

of infection 

Organs examined 

Rat number 































Bone marrow 





Heart _ 




Liver _->« 










Days killed after infection 

* Mouse 

Since, the animals used in the above experiment w^re killed 
by bleeding, the results obtained are to a certain degree ques- 
tionable. To rule out the possible influence of bleeding in 
the natural distribution of the parasites in the various organs, 
another series of rats was infected at various intervals and 
killed with chloroform inhalation. The impression smears 
of these animals were made, stained and examined the day they 
were prepared. The majority of the rats used, were killed at 
the time the tail blood was swarming with trypanosomes (3+ 
to 4-4- per microscopic field) as shown in Table 3. A perusal 
of the results presented in the above cited table one will note 
that the lung, liver, heart and kidneys grouped together bear 
an almost identical trypanosome burden per microscopic field 
as that of the tail blood, the range being 3-+- to 4+ for both. 
Comparing the above organs with the rest, there is a decided 
heavier concentration of the organisms in the lungs, liver, heart 
and kidney than in the brain, spleen, bone marrow and lymph 
glands (rats 2 to 5 and 11 to 20). A comparative study of 
tables 2 and 3 will show that no such decided change in the 
intensity distribution of the trypanosome has taken place in 
the lungs, liver, heart and kidneys as a result of bleeding be- 
yond a slight degree in the kidneys and lungs. However, in 
the case of the bone marrow, brain and spleen the change in the 

Yutuc and Sher: Distribution of Trypanosoma evansi 161 

density is quite marked. In some of the rats that were killed 
when the tail blood was lightly infected with trypanosomes 
(Table 3, rats Nos. 1, 2, 8, 9, and 10) the parasite burden 
of the peripheral blood is almost the same as that in the heart, 
liver, lungs and kidneys. Indirectly, these five rats serve to 
bridge the gap between the prepatent period and the patent 
period of maximum trypanosome density in the peripheral 
blood. Again the brain, spleen and bone marrow carry a cor- 
respondingly lighter trypanosome burden than the peripheral 

The above data present an added weight to the statement 
that the more blood the organ contains the more parasites it 
harbors. The liver, heart, kidneys and lungs being more vas- 
cular than the rest of the organs studied, therefore, presented 
the heaviest trypanosome density at least in an animal \^here 
the resistance is low as in the case of rats. The density of 
the organisms in the peripheral blood is almost identical with 
that in the liver, heart, kidneys and lungs but decidedly heavier 
than the trypanosome density of the spleen, brain, bone mar- 
row and lymph glands. In this case, it is erroneous to sub- 
scribe fully to the statement that the parasites are evenly dis- 
tributed in the various organs of the body. Further, the 
low density of the trypanosomes in the bone marrow, spleen, 
and lymph tissues may be attributed to the statement of Mar- 
tini (1903) who states that these organs, constituting the 
great bulk of the reticulo-endothelial systems, are quite active 
in the destruction of the organisms. 

Table 3. — Distribution of trypanosomes in the body of rats during the 

patent period 

Rat number 

Organs examined 








2 + 
2 + 

















2 + 


Bone marrow _ 

Brain - 



Kidnevs -- 


Spleen -.- 

Lymph gland-, 

Tail blood 






2 + 











} killed after infection 


The Philippine Journal of Science 


Table 3. — Distribution of trypanosomes in the body of rats during the 
patent period — Continued 

Organ? examined 

Rat number 











Bone marrow 























Heart .._ 



Kidneys . ! 


Lnng^ . ..... 



Lymph gland 

Tail blood 





3 + 














Days killed after infection 

Subpatent period. — In the subpatent period study, guinea 
pigs were killed as early as fourteen days and as late as one 
hundred and eighty-four days after infective inoculation of 
surra blood (Table 4). The picture of trypanosome distri- 
bution is quite different from the rats. Relative to the fre- 
quency of the microscopic finding of the trypanosome in the 
organs examined, the brain, and the lymph glands show the 
parasites eight times out of the fifteen guinea pigs thus far 
employed in the investigation. The spinal cord comes next 
with four positive results out of the numbers used ; kidney and 
liver 3 times and the bone marrow and spleen 2 times; and 
in the lungs and heart the organism was only noted once. In 
the adrenals the trypanosome was noted two times in the six 
cases that this organ was examined. In regards to the densi- 
ty, although quite variable in the same organ of the differ- 
ent animals of the species, the brain tissue demonstrated the 
heaviest density of trypanosome its maximum being S-f and 
the minimum +. Taking the animals as a whole it will be 
noted that the organism was noted in all organs examined. 
For comparative study a number of guinea pigs with positive 
peripheral blood ( + ) were killed and distribution studied. 
The brain and lymph glands showed the heaviest density, the 
former being 3-f-, the latter 2+. The rest of the organs gave 
almost the same intensity as the surface blood. In discussing 
the results of this series, at least in the organs consistently 
used, the general statement '*the more vascular the organ, the 
heavier is the trypanosome density and that the trypanosome 

Yutuc and Sher: Distribution of Trypanosoma evansi 163 

density of the peripheral blood is directly proportional to the 
density of the organisms of the vascular organs given in the 
rats" does not seem to fit in with the results obtained in the 
guinea pigs. Further, the brain, lymph glands and the spinal 
cord are certainly less vascular than the heart, liver, kidneys 
and lungs, yet the burden of trypanosomes is significantly 
heavier in the former than in the latter. In the guinea pig the 
organism has the strongest affinity to settle in greater number 
in the nervous tissue than the rest of the organs both in pat- 
ent and subpatent periods. To elucidate the different find- 
ings between the rats on one hand and the guinea pigs on the 
other, one has to look into the differences in the resistance of 
the two animals to surra infection and the course of the disease 
when the animals were killed. In this particular instance the 
rat represents a susceptible animal whereas the guinea pig a 
resistant subject. In fact, Tubangui and Yutuc (1931) stated 
that a varying degree of resistance is exhibited by different 
species of surra animals. This is indicated by their ability 
to keep in abeyance for a shorter or a longer period of time the 
rapid and permanent proliferation of the surra trypanosomes. 
Thus, on the one hand the rat possesses a low type of resist- 
ance for, in this animal the infection runs an acute progressive 
course, the parasite undergoing a constant increase in num- 
ber and death taking place when the number of the parasites 
in the blood has reached a miximum level. The guinea pig, on 
the other hand, possesses a high type of resistance, for in 
this animal the infection follows a chronic relapsing course and 
death usually occurs when the parasites are scarce in the cir- 
culation. In vitro study of the sera of rats and guinea pigs 
infected with surra organisms Kligler (1931) demonstrated 
trypanolysin in the guinea pigs but not in rats. In the latter 
no lysin existed at any stage of trypanosome infection. As a 
result of splenectomy studies in rats, the main defense has been 
found to be a cellular one in this species. These lytic bodies 
in the guinea pig and other resistant species have also been 
repeatedly demonstrated by a number of investigators in vivo 
studies (Diesing, 1908; Kleine and Hollers, 1906; Taliaferro, 
and Johnson, 1926). It is probable that this resistance m 
the guinea pig induced the trypanosome to localize in less vas- 
cular organs such as the brain, spinal cord and the lymph 
glands when the peripheral blood was negative in microscopic 


The Philippine Journal of Science 


Tabije 4. — Distribution of surra trypanosomes in the body of guinea pigs 
during subpatent period 

Organs examined 


pig number 


























Bone marrow 

















Lymph glands 



Spleen I./ 

Spinal cord 
















srs killed 




An inquiry into the distribution of Trypanosoma evansi 
(Steel, 1885) in the body of rats and guinea pigs was 
performed. It has been found that in rats during the prepat- 
ent period the organisms are scarce in the internal organs 
such as the bone marrow, brain, heart, kidneys, liver, lungs, 
spleen and lymph glands. No indication tending to suggest 
that the organisms multiply and accumulate in considerable 
number in the organs examined during the initial period was 
found. During the patent period there seems to be a direct re- 
lationship between the density of the trypanosomes in the pe- 
ripheral blood and in the heart, liver, lungs, and kidneys. In 
the spleen, brain, bone marrow and lymph glands, the distri- 
bution intensity is lower than in the peripheral blood. While 
bleeding has little or no effect on the distribution of the or- 
ganisms in the liver, heart, lungs and kidneys, a significant 
decrease in the bone marrow, brain and spleen was recorded. 
During the subpatent period in guinea pigs the brain, lymph 
glands and the spinal cord which have light trypanosome bur- 
dens during patent period, presented a comparatively heavier 
trypanosomes density. The resistance of the animal to the in- 
fection may possibly be the factor responsible for this finding. 
The statement that the organisms are evenly distributed in 
the different organs of the body was not sustained and that the 
multiplication of the trypanosome takes place in the body 
fluids seems justified. 

Yutuc and Sher: Distribution of Trypanosoma evansi 165 


Craig, C. F. and E. C. Faust. Clinical Parasitology. Phila., Lea and 

Febiger, 1937. 3-733 pp. 
DiESiNG, Ein Immunisierungsversuck gagen die Tsetse krankheit der 
Kinder in Kamerun. Arch. f. Schiffs— u. Trop. Hyg. 9 (1908) 
Klbinb, F. K. and B. Mollers. Ein fur T. brucei specifiches Serum 
und seine Einwirkung auf T. gambiense. Ztschr. f. Hyg. u. In- 
fektionskr. 52 (1906) 229-237. 
Kligler, I. J. Susceptibility and resistance to trypanosome infection. 
VIII: In vitro demonstral;ion of specific agglutinating and trypa- 
nolytic antibodies in the serum of infected guinea pigs. Ann. Trop. 
Med. and Parasitol. 25 (1981) 377-393. 
Manrbsa, M. and B. M. Gonzalez. Studies of surra? IV. Variability 
in size of the trjrpanosomes and density of infection, natural and 
acquired immunity. Philip. Agric. 24 (1936) 716-751. 
Martini, Erich. Ueber die Entwickelung der Tsetseparasiten in Sa- 

ugethieren. Ztschr. f. Hyg. 42 (1903) 341-350. 
Mtjsgrave, W. E. and M. T. CiiEGG. Trypanosoma and trypanosomiasis, 
with special reference to surra in the Philippine Islands. Bur, Govt. 
Lab. No. 5. (190B) 4-248. 
Plimmer, H. G. and R. J. Bradford. A preliminary note on the mor- 
phology and distribution of organism found in the tsetse-fly disease. 
Proc. Roy. Soc. 65 (1899) 274-281. 
Taliaferro, W. H. and T. L. Johnson. Zone phenomena in vivo trypa- 
nolysis and the therapeutic value of trypanolytic sera. Jour. Proc. 
Med. 1 (1926) 85-123. ^^ ^ 

Tubangui, M. a. and L. M. Yutuc. The resistance and the blood sugar 
of animals infected with T. evansL Philip. Jour. Sci. 45 (1931) 93-107. 
Schilling. Bericht ueber die Surra— -krankheit der Pferde. Centralbl. 

f. Bakt. 7, Abt. 30 (1901) 545-551. „ , ^. 

VoGES, O. Das Mai de Caderas der Pferde in Sudamerika. Berl. tie- 

rarztl. Wehnschr. Oct. 3 (1901) 597-598. 
Yutuc, L. M. Experimentar Studies in the curative ^^^^tment of surra 
in native horses in the Philippine Islands. Philip. Jour. Sci. 54 (1934) 


By Moricb Vanoverbergh 
Belgian Missionary, Bauko, Mountain Province, Philippines 



A great many adverbs are expressed in Iloko by simple 
adjectives ; by substantives, adjectives, and so on, in the oblique ; 
by verbs with an adverbial meaning; and by substantives, verbs, 
and so on, with adverbial prefixes, infixes or suffixes, or with 
adverbial reduplications. 

Many of these forms are found scattered about in former 
publications, under different headings. To be more or less 
complete and to supply as much as possible all omissions, I 
shall give the following list, which may serve at the same time 
as a recapitulation. Later on I shall study some of the most 
used adverbs and adverbial expressions in particular. 

I. Adjectives used as adverbs. Examples: 

nadards ti parmagndna 
naindyad ti patmggundyyo 
napintds ti panagabelda 
naalds ti panagsaoda 
nalaing ti panagsaritam 
nagagH nga agsursHro 
nasadutkayo nga agsHrat 

na'&yong a mangisilro 

he walks quickly. 

you move slowly. 

they weave nicely. 

they speak unbecomingly. 

you speak fluently. 

he studies diligently. 

you are lazy writers (you write 

lazily) . 
he is a severe teacher (he teaches 

severely) . 

II. The oblique taken adverbially. Examples: 

mapdnkayo iti rumabii 
mapdn iti masansdn 
immdykami iti rabii 
agsublikayo iti bigdt 
mangdn iti mabaydg 
madHsadanto iti agnandyon 
simmangpet iti ngdlay ti rabii 
pumdnawkami iti apagbiit 
napdn iti snnoin daydi a Idwas 
iti naminsdn nga alddw 
iti naminsdn a panangrabiida 

go at nightfall. 

he often goes. 

we came in the evening. 

come back in the morning. 

he eats long. 

they will be punished eternally. 

he arrived at midnight. 

we go away for a while. 

he went the following week. 

on a certain day. 

once, while they had supper. 



The Philippine Journal of Science 


addd iti un4g 

matiirogkami iti pagbaetdnda 
agnaedkayo iti tengngdna 
makisaritada iti rudr 
nagydn iti nagsengngatdnda 
immuli iti ngdtona 
napdn iti babdna 
addd iti makatigidna 
agsdo iti naalds 
magndka iti nadards 
nagsakdy iti namindud 
agsaoda iti adu 

he is inside. 

we sleep between them. 

you live in its midst. 

they talk outside. 

he was located between them. 

he climbed above. 

he went below. 

it is at his left. 

he speaks unbecomingly. 

walk quickly. 

he rode twice. 

they talk much. 

III. Verbs with an adverbial meaning. Examples : 

sansdnem ti agludlo 
bayaginda ti agsmita 
mndpami ti nagbantdy 

agdamd a mangdn 
iniwyadmo ti nagnd 
nagindyadda a nagtardy 

in-indtenda ti pumidut 

dina met binaydg a pinagkot- 

pray often. 

they talk long. 

we watched early. 

he is in the middle. 

they go out, 

it goes up. 

it goes down. 

he is eating just now, 

you walked slowly. 

they ran slowly. 

be quick. 

they pick them up little by little. 

we go fast. 

he was not long in getting thin. 

IV. Adverbial prefixes, infixes and suffixes: 

1. Notion of concomitance: 

a. The Prefix ka and the combination ka . . . an. Cfr, The 
Substantive, III. Formation of Substantives, III, 4; The Ad- 
jectives, III. Special Forms. 1; and The verb. The Adjectival 
Prefix maki. b. II. and so on.^ 

b. The Prefix maki. Cfr. The Verb, The Adjectival Prefix 

c. The Prefix si. Cfr. The Adjective, III. Special Forms. 7; 
The Verb, The Adjectival Prefix ag. b. Ill, A. 4 ; The Adjectival 
Prefix ma. h. 1. and so on.' 

lAnthropos 26 (1931) 472-478, 480-481; Philip. Jour. Sci. 69 (1939) 
* Philip. Jour. Sci. 69 (1939) 235-237. 
•Anthropos 26 (1931) 484-485; Philip. Jour. Sci. 69 (1939) 231, 239. 

Vanoverbergh: Iloko Adverbs and Prepositions 169 

d. The Prefix tagi. Cfr. The Verb, The Adjectival Prefix 
ag. b- III. B. 1 ; The Substantial Suffix en. b. IL 1. and so on.* 

e. The Prefixes sanga and sangka combined with Prefix i 
Cfr. The Verb, The Substantial Prefix i. b. II. Note 6. B.^ 

2. Notion of similarity : 

a. The Prefix sinan or sinin and mara. Cfr. The Adjective, 
III. Special Forms. 3; and The Verb, The Adjectival Prefix 
ag. b. III. A. 3.^ 

b. The Suffix en and the combination pang ... en. Cfr. 
The Substantive, III. Formation of Substantives. V.*^ 

3. Notion of reciprocity : The infix inn. Cfr. The Substantive. 
III. Formation of Substantives. III. 6; The Verb, The Prefix 
ag. h. III. A. 2; The Prefix maki. b. II. 2. and so on.^ 

4. Notion of recency: The prefixes fca, apag and kapag. 
Cfr. The Adjective, III. Special Forms. 2.» 

5. Notion of repetition: 

a. The Infix an. Cfr. The Verb, The Adjectival Infix an, 
and so on, ^^ 

b. The Prefix kara. Cfr. The Verb, Prefix kara; Suffix en; 
Prefix I B. I. 3.11 

6. Notion of possibility: The Prefixes maka and ma. Cfr. 
The Verb, Prefix maka; Prefix ma. (adjectival and substantival), 
and so on. ^^ 

7. Notion of instrumentality: 

a. The Prefix pag, and so on, and the Prefix pa. Cfr. The 
Substantive, III. Formation of Substantives. VI ; The Verb, Sub- 
stantives with Verbal Notions, and so on. ^^ 

b. The Prefix i. Cfr. The Verb, Prefix i.^^ 

8. Notion of being of less importance: The Infix urn. Cfr. 
The Verb, The Infix um; Verbs Implying Order or Permission, 
pa ... en ^^ 

* Philip, Jour. Sci. 69 (1939) 231; 71 (1940) 12. 

"Philip, Jour. Sci. 71 (1940) 20. 

•Anthropos 26 (1931) 482-483; Philip. Jour. Sci. 69 (1939) 231 

^Anthropos 26 (1931) 474-475. 

sAnthropos 26 (1931) 473-474; Philip. Jour. Sci. 69 (1939) 230^231, 

^Anthropos 26 (1931) 481-482, 
^'^ Philip. Jour. Sci. 69 (1939) 241-242. 

"Philip. Jour. Sci. 69 (1939) 240-241; 71 (1940) ^\]lf^;^^ ^^^ 
-Philip. Jour. Sci. 69 (1939) 233-235, 239-240; 72 (1940) 429-440. 
-Anthropos 26 (1931) 475; Philip. Jour. Sci. 69 (1939) 247-249. 
"Philip. Jour. Sci. 71 (1940) 15-20. 
t^ Philip. Jour. Sci. 69 (1939) 232-233, 71 (1940) 30-31. 

170 The Philippine Journal of Science ^^^ 

9* Notion of quantity or quality: The Prefixes nag, naka, 
sunmn^ka, and the Combinations nai • . • an and nain , • . 
an, Cfr. The Adjectives, II. Comparatives and Superlatives, 
passim, ^^ 

10. Pretending to be so: The complex prefix agin. Cfr. The 
Verb, Prefix ag. b. III. B. 2. and so on." 

11. Almost: The combinations of the prefixes tagi, tan, 
art, and aHn with the suffixes an and en. Cfr. The Verb, The 
Adjectival Combinations with tari, etc. ^^ 

12. Locative: 

a. The Suffix an, Cfr. The Substantive, III. Formation of 
Substantives, III. 3; IV; The Verb, Substantive with Verbal 
Notions; The Substantival Suffix an\ The Prefix ma. II. Note 
18. a. and so on, ^^ 

b. The Prefixes i and ipa. Cfr. The Vei^b, Prefix i\ Prefix 
ipa, and so on. ^^ 

c. The combination pakin. ... en. Cfr. The Verb, Prefix 
ipa. A. 3. Note 7.^1 

d. The terms addd and awdn. Cfr. To have and to heP 

13. Frequentative : Cfr. The Verb, Frequentative Prefixes.-^ 

14. How and when, where, why and to whom: Cfr. The 
Verb, Substantive with Verbal Notions,^^ 

15. Ordinal numbers, multiplicatives, distributives, indefinite 
numbers, fractional numbers: 

a, Cfr. The Number, II, III, IV, and V; The Verb, Suffix 
en. b. II. 3. and so on. 25 

b. The combination pi . . . en ov pin ... en and the Pre- 
fixes ipi or ipin and ikapi or ikapin. Cfr. The Verb Combina- 
tion pi . , . en, and so on.2« 

16. Reaching a certain part of the body : The Prefixes aga and 
paga. Cfr. The Verb, The Adjectival Prefix aga. ^'^ 

i«Anthropos 26 (1931) 477-479. 
"l>hilip. Jour. Sci. 69 (1939) 231-232. 
"Philip. Jour. Sci. 69 (1939) 245. 

i^Anthropos 62 (1931) 472, 474; Philip. Jour. Sci. 69 (1939) 247, 
250-253; 71 (1940) 12-15; 72 (1940) 439. 
^Philip. Jour. Sci. 71 (1940) 15-22. 
^Philip. Jour. Sci. 71 (1938) 21. 
** Philip. Jour. Sci. 66 (1938) 417-431. 
« Philip. Jour .Sci. 69 (1939) 245-247. 
^Philip. Jour. Sci. m (1939) 247-248, 250-255. 
*Anthropos 28 (1933) 714-720; Philip. Jour. Sci. 71 (1940) 12. 
^Philip. Jour. Sci. 71 (1940) 23-24. 
^ Philip. Jour. Sci. 69 (1939) 242-243. 

Vanoverbergh: Iloko Adverbs and Prepositions 171 

V. Adverbial reduplications: 

1. Notion of easiness, and so on: reduplication of the first 
open syllable. Cfr. The Verb, Reduplication of Verbal Stems, 
passim. 28 

2. Notion of reciprocity, rivalry, repetition, and so on: re- 
duplication of two or more syllables. Cfr. The Substantive, III. 
Formation of Substantives. Ill, 6; The Verb, Prefix ag, h. III. 
A. 2, Prefix maki. b. II. 2; Reduplication of Verbal Stems, 
passim. ^^ 

3. Notion of similarity and so on: reduplication of the first 
open syllable with the next consonant if there is any. Cfr. The 
Adjective, Special Forms. 3; The Verb, Prefix a^r. b. III. A. 
3 ; and so on. ^^ 


I. 1. The ordinary simple adverbs of time are: 

a. For the present: itd and itattd (reduplication form, the 
initial i being replaced by the ligature t). Examples: 

adddda itd this time they are here. 

sadin ti papandnyo itd where do you go at this time? 

inkayo itattd go now directly. 

inkami mangdyo itattd we go to gather wood presently 

b. For the past: implying a near past: itdy and itattdy 
reduplicated form); implying a more or less distant past: 
idi. Examples : 

immdy itdy he came just now 

nagsangit itdy he cried just now. 

adddda itattdy they were here a short while ago, 

sangsangpitna itattdy he has just come home a little while 


natdy idi he was dead at the time. 

pimmdnaw idi he was gone then. 

nagsakitak idit ta then I got sick there. 

Note 1. In ordinary conversation, when itdy and idi follow another 
word, their initial i is very often replaced by the ligature t, especially 
when the other word ends in a vowel. Examples: 

Sikdt tay (for: Sika itdy) it was you just now. 

isMdi (for: isH idi) it was he at the time. 

» Philip. Jour. Sci. 75 (1941) 207-211. 

«»Anthropos 26 (1931) 473-474; Philip. Jour. Sci. 69 (1939) 230-231, 
236^237; 75 (1941) 207-211. ^_^^^ 

aoAnthropos 26 (1931) 482; Philip. Jour. Sci. 69 (1939) 231; 75 (1941) 


172 The Philippine Journal of Science ^^^^ 

2. Complex adverbs of time are formed by the addition of 
a substantive (either alone or accompanied by a qualifying 
word), an adjective, and so on, to: 

a. The demonstrative itoy or itd (preceding, ligature a or 
nga), for the present Examples: 

itoy nga alddw today. 

itoy a bigdt this morning. 

itoy a bulan this month. 

itd nga alddw today. 

itd a malim this afternoon. 

itd a rabii this evening. 

itd a nasdpa now as it is early. 

itd a nalddaw now as it is late 

Note 2. Itoy may follow immediately, but then the article precedes 
the substantive, and so on. Examples: 

itdy alddw itoy today. 

b. The demonstrative itdy or idij rarely idiay (no ligature), 
for the past. Examples: 

itdy agsdpa early this morning. 

itdy maUm this afternoon. 

itdy kud a while ago. 

itdy parbdngon this morning very early. 

itdy napdn a tawen last year, 

itdy kallabes a tawen last year. 

idi kalmdn yesterday. 

idi rabii last night. 

idi malim kalmdn yesterday afternoon. 

idi bigdt kalmdn yesterday morning. 

idi kallabia ti talld a bulan three months ago. 

idi %apdn a tawen last year. 

idi maikatlo a tawen three years ago. 

idi kud then. 

idi nabaydg long ago. 

idi naminsdn nga alddw the day before yesterday. 

idi (ka) sangaalddw the day before last. 

idi kasangatawen the year before last. 

idi katimmawin three years ago. 

ididy nakabutbuteng nga alddw on that terrible day. 

c. The conjunction intono, no, tono (ordinary forms) , attono, 
antono (rarely used), for the future. Examples: 

intono bigdt tomorrow. 

bigdt no bigdt tomorrow morning. 

maMm no bigdt tomorrow afternoon. 

tono mxilem this afternoon. 

Vanoverbergh: Iloko Adverbs and Prepositions I73 

attono alddw 

antono rabii 

no rabii 

(in)t6no kud 

intono mabaydg 

intono umdy a bulan 

intono maikapdt a bulan 

intono maldbes ti uppdt a bulan 

intono arinunos ti bulan 

intono sangatawen 

intono sangaalddw 

intono sangaalddw a maysd 

later on in the day. 

this evening. 

this evening. 


much later. 

next months. 

after four month. 

after four months. 

at the end of the month. 

after two years. 

the day after tomorrow. 

after three days. 

Note 3. When these adverbs of time follow the verb, no ligature con- 
nects the two. Examples: 

umdyka Hoy nga aldaw 
agdladkami itoy nga alddw 
napdnda itdy alddw 
sinuronnakami itdy agsdpa 
kasangsangpetda itdy malem 

nagtdlawda idi rabii 
inddalko idi nabaydg 
immdy idi alddw kalmdn 
mapdnto no bigdt 
matdyto no mabiit 
mapdnkami intono agsdpa 
umdyak no kud 
sumangpet intono kud 

come today 

we make a fence today, 
they went about midday, 
he teased us early this morning, 
they just came home this after- 
they escaped last night. 
I learned it long ago. 
he came yesterday about midday, 
he will go tomorrow 
he will die soon, 
we shall go early tomorrow. 
I shall come later, 
he will arrive later. 

Note 4. The term agdanm may be used to indicate the present; when 
used in the past (nagdamd)^ it indicates concurrence of events. Ex- 
amples : 

agdamd a mangdn 
agdamdda nga agsurstirat 
nxigdamdda a nagsdngit idi 
iti agdamd a bulan 
iti agdamd a tawSn 
iti tawen nga agdamd 

he is eating presently, 
they are writing just now. 
they were crying at the time, 
the current month, 
the current year, 
the current year. 

Note 5. The combination ka , , . an (past kina . . . ati) adds the notion 
of "the following'* to the substantive which indicate the period of time. 

iti kabigatdnna 

idi karabiidn ti isasangpetko 

intono kabigatdnna 

idi kinabigatdn ti daydi alddw 

the next day. 

the evening after I came home. 

the next day (future). 

the next day (past). 

174 The Philippine Journal of Science ^^^^ 

II. To indicate that something is done every day, every month, 
and so on: 

1. The infix in is joined to the stem. Examples: 

inalddw every day. 

hinigdt every morning. 

rinahii every evening. 

umdy iti binuldn he conies every month. 

tinaw4n ti panagsakitko I am ill every year. 

2. The term patindyon or (kan) kandyon is connected with the 
noun by a or 'iiga. Examples : 

patindyon nga alddw every day 

patindyon a hMan every month. 

kandyon a higdt every morning. 

kankandyon a malem every afternoon. 

Note 6. When the infix in is joined to the stem and its first two 
syllables are reduplicated besides, the recurrence of the event is emphasized. 
Examples : 

inaldaw-alddw day by day. 

binigatbigdt every morning without fail. 

rinabirabii every evening without fail. 

tinawentawin every year without fail. 

III. The prefix maka (past: naka) joined to the substan- 
tive which indicates the period of time, means that something 
lasts once the duration implied by the substantive. If the 
duration is longer, the Iloko connect the number with the 
substantive, followed by a possessive pronoun, by the ligature 
a or nga. Examples : 

makab'ulankayonto ididy you will stay there a month. 

nakatawSnda ididy babd they stayed a year in the lowlands. 

dud nga alddwmi ididy we stayed there two days. 

dud pulo nga alddwtayo ididy we stayed twenty days in the 

kabakiran forest. 

sangapMo a tawinmi ididy we stayed there ten years. 

Note 7. Agpatndg (past: Tiagpatndg), from the complex prefix agpa 
and the stem tinndg, means: the whole night; agmalmalem (past: nag- 
malmalem) means: the whole day. Examples: 

agpatndg nga agsursiHrat he passes the night writing. 

nagpatndgkayo a nagsugsugdl you spent the whole night gam- 
agmalmalemda nga agsangsa- they weep the whole day. 

nagmalmaUmkaim a nagnagnd we walked the whole day. 

Note 8. Sometimes the prefixes maka and makanika (past: naka and 
nakanika) are joined to the cardinal numbers in the same way as the prefix 
maikd (of ordinal numbers), to indicate a period of days. The prefix maka 

Vanoverbergh: Iloko Adverbs and Prepositions 175 

may also be joined to the distributives with the same meaning. "One day" 
is makaysa (past: nakaysa). The corresponding interrogatives are: wa- 
kamano and makasagmamano. If the period is not a number of days, 
but a number of months, and so on, this must be expressed in the sentence. 
These forms, however, are used very rarely. Examples: 

makatlodanto nga agydn ididy they will stay there three days. 
makamamkayo ditoy how many days do you stay here? 

makanikatlo pulokami ket dud we shall stay twenty-two days. 
makaysddanto they will stay one day. 

makapdtda ididy they stay there four days. 

makasagwawalokamto we shall stay eight days. 

makapdt a tawen ditoy he stays here four years. 

Note 9. MakamanS and makamanmano (rarely used) mean: a long 
time, much time, difficult. Examples: 

sadn a makamano daytd nga that work is easy. 

makamanmanoka a baonen it is very hard to send you some- 


IV. A peculiar adverb of time is en which is often joined 
to the last word of the sentence and means: already, now, 
and so on. We saw one of its many uses when treating the 
comparatives and superlatives of adjectives,^! and we shall 
see another one in the disintegration of the adverb manen 
"again.*' 32 cfr. also Phonology, VI Syncopations. II. B. c. and 
IV. B. 1.^ Examples: 

adddn here he is now. 

umdykan come along now. 

simmangpiten he has arrived already. 

nakaddnon ti lakdyen the old man got there already. 

sidkon it is my turn now. 

aldemon all right, take it now. 

maydtakon I am willling now. 

kudkon now it is mine. 

kitdem ti baketen look at the old woman now. 

addd mettin liere he is now though. 

madikamin we |on't want it any more. 

addd nga agtardyen there he is running now. 

nagtdlawsa met ti dsomon your dog escaped already, didn t 


inaydtnan s^e loved him already. 

madamdn ^^ ^^ at it. 

idi kudu and then at the time- 


arise now. 

^^Anthropos 26 (1931) 479. 

»^See below: VI. Adverbs of Quality and Manner, IV. It^. 

«^Anthropos 23 (1928) 1936, 1940. 

8104 4 


The Philippine Jourtml of Science 


manggapon nga agbanndwag 
anidn, daytoyen 


kondk Idbn nalaklakdda 


sadn ngaminen ayd nga agsur- 

ania ketdin ayd 

it begins to dawn already, 
what now, what is the matter with 

this one? 
stop it now; it is enough now, 
I thought though that they were 

there is no more; he is not here 

any more; it is done for. 
well, indeed, is he not writing any 

more now? 
stop it, what is the matter there? 

what can be done? (resignation) 

V. Other adverbs of time: 
1. Patindyon, {kan) kandyon and agnandyon: 


agludlo a patindyon 
patindyonda nga agsdngit 
patindyon nga agrabrdbak 
kankmndyon a tumardy 
agbasbdsa a kankandyon 
kankanyon Ideng a nagtalinaed- 

da iti kappid 
kandyon nga agddwat 
kastd a kandyon 
naragsdkda nga agrmndyon 
madusada iti agnandyon 

he always prays, 
they always weep, 
he is always jesting, 
he always runs, 
he is always reading. 

they always remained 4n peace, 
he always asks for things, 
it is always so. 
they are always full of Joy. 
they are punished eternally. 

2. Masansdn: (past: nasansdn), rmmin-adu (past: mtmin- 
adu and nasarantd: "often/* Examples : 

masansdn nga umdy 
masansdn a sasaoinvm 
mxisansdn ti panagkatdwana 
agsdngit iti masansdn 
agladingit a masansdn 
mamin-adik nga agbartek 
namin-adA a nagunget^ 
kimmagdt iti namin-adu 
nasardnta ti panagddwatna 

3, No dadduma: "sometimes" 

imm4y no daddiiima 
no dadduma agsdngit 
mxingdnkami no dadduma iti ti- 

no dadd4m4i napdnkami ididy 


he often comes. 

he often says so. 

he often laughs. 

he often weeps. 

he often grieves. 

he often gets drunk. 

he often was angry. 

he often bit. 

he often asks for things. 

(no ligature). Examples: 

sometimes he came, 
sometimes he weeps, 
sometimes we eat bread, 

sometimes we went to Bangar. 

Vanoverbergh: Iloko Adverbs and Prepositions 177 

4. Sagpaminsdn: '*now and then." Examples: 

mangdlakami iti sagpaminsdn we take some now and then. 

umdy no sagpaminsdn he comes now and then. 

sagpaminsdin ti yadymi we come now and then. 

sagpamifisdn nga agbartek he gets drunk now and then. 

5. (Pas) pasardy: "occasionally" (preceding)* Examples: 

paspasardy umdy he comes occasionally. 

pasardy naghartek he got drunk occasionally. 

pasardyda tumalidw they look back occasionally. 

6. Ifganngdni and dandani: "nearly, almost", and so on 
(preceding, no ligature). Examples: 

ngangdnika sapllten I shall soon whip you. 

tiganngdni rabiien it is almost evening already. 

dandani pumdnaw he is near going. 

dandani a las dos it is nearly two o'clock. 

dandani ni ulitegko itdn my uncle is almost here now. 

7. Madamdnma: "soon." Examples: 

sumangpH no madamdamd he will arrive soon. 

no madamdamd umdyak I shall come soon. 

iti madamdamd bassit a little later. 

8. Biit and bassit: entreating; "literally for a moment" 
(following, no ligature). Examples: 

swmrekka biit come in for a moment. 

inka bassit ididy Sdnta go once to Santa. 

i(nka bassit alden ti suratko please, go to take my letters. 

I. The three simple adverbs of place, corresponding to the 
three demonstratives, are: 

Complete form Abbreviated form Corresponds to 

(here) toy daytoy (this) 

ditd (there) ta daytd (that) 

ididy, sadidy (there) didy daydidy (that) 

adddkam ditoy we are here. 

simmangpet ditoy he arrived here. 

madi a mangdn ditoy he won't eat here. 

addd masakit ditoy is there anyone sick here? 

nangasdwa ditoy he married here. 

ditoy nakataydnna he died here. 

adii ti bagds ditoy baldymi there is much rice in our house 


ti kiikua ditoy lubong the earthly goods. 

umdykat toy come here. 


The Philippine Journal of Science 


awdn toy ti kaydtko 

isu ti adddt toy 

awdn ti maydt toy 

isH ti nakiasdwa kenkudnat toy 

awdn ditd 

madida dmin ditd 

addd nangdla ditd 

ditd ti papandnda 

adH ti bagds ditd haldyyo 

awdn ditd ydnmo 
inkat ta 

adddt ta ti birokek 
umdyaktot ta no umimhdgak 
no umdykamit ta agungStka 
nakaddnondat ta 
nagsakitak idit ta 
napdnkami ididy 
Madida a matHrog ididy 
ada ti bagds ididy 
inkamto gumdtang ididy 
dagiti ubbing ti napdn ididy 
ididy ti nakasardkatnmi 
inkat didy 
addddat didy 
bassit ti maydt kanot didy 

isudat didy 

adddnsat didy ni Luis 

what I want is not here. 

it is he who is here. 

nobody is willing here. 

she married him here. 

it is not there. 

do they all refuse there? 

someone took it there. 

they go there. 

is there much rice in your house 

it is not at your place there, 
go there. 

what I look for is there. 
I shall come there when I get well, 
when we come there you are angry, 
they got there, 
then I got ill there, 
we went there. 

they do not want to sleep there, 
there is much rice there, 
we shall go to buy some there, 
the children went there, 
we met him there, 
go there, 
they are there, 
they say that few people there are 

there they are. 
Lewis is there, I believe. 

Note 10. There is no difference between the use of the complete forms 
and that of the abbreviated ones, and, when these adverbs follow the 
verb and so on, (except an occasional t) no ligature is used to connect 
them all this may be seen in the above examples. 

Note 11. Sadidy refers to a place which has been alluded to before. 
Examples : 

inka didy simbadn sadak ura- go to the church and wait for me 

yen sadidy there. 

napandk ididy Dingrds ket sa- I went to Dingras and met him 

didy ti nakakitdak kenkudna there. 

Note 12. In some districts the Iloko use sadi instead of idiay^. Ex- 

Sadin ti papandm? — Sadi where do you go? — There. 

Note 13. Sometimes nay is used as ^n adverb of place, but it is not 
very polite term. Examples: 

nay aldem 

ditd nay lansdngan 

bumalldsiwka ditd nay dlad 

here, take it. 
there in the street, 
cross that fence there. 

^See below: The Preposition, VI. 

Vanoverbergh: Iloko Adverbs and Prepositions 179 

Note 14. Ni may be added to words indicating place or location, and 
sometimes also to other words. It more or less represents the English 
"see." Examples: 

adddt toy ni 
kitdem ni 
awdn ni 
kdstoy ni 

it is here, see. 
look here, 
it is not here, see. 
like this, look. 

Note 15. Ditoy may be used as a substantive including the notion of 
an adverb of place. Examples: 

nasakit ti ditoyko 
addd gaddil iti ditoyna 

this hurts (showing where), 
he has itch here (showing where). 

11. The following adverbial expressions sometimes occur: 

adtoy or atWy 

addaytoy (rarely used) 






Examples : 

adtoy ti adipenmo 
adtoy ni apo Juan 
adtoy a hinaon ti lakdy 
adtoy ti ydnna 
addaytoy ti ubing 
addaytd ti birokem 
addaydidy ti asinyo 
addagitoy kami 
addagididy ti dsoyo 

Contraction : 
Contraction : 
Contraction : 
Contraction : 
Contraction : 

addd and toy 

addd and daytoy 

addd and daytd 

addd and daydidy 

addd and dagitoy 

addd and dagitd 
addd and dagididy 

behold thy servant. 

here is Mr. John. 

here he is whom the old man sent. 

here he is. 

here is the child. 

there is what you look for. 

there is your salt. 

here we are. ^ 

there they are now. 

there are your dogs. 

III. To indicate the relative position of two or more objects, 
the prefixes akin or makin (past: nakin) are joined to the stem 
implying the position of one of the objects. Cf r. The Adjective, 
III. Special Forms. 4; The Verb, The Prefix ipa. A. II. 2. 
Note 7.35 Examples : 

akinddya daytoy 
daydidy ti akinrabdw 
makinkanawdn ti ydnna 
isu ti nakinkatigid 
ipdnmo iti makinldud 
ti makin-uneg 
ti makinrudr a kdyo 
addd iti makinngato 

this is on the east side, 
that is on the upper side, 
he stands at the right, 
he was at the left, 
carry it to the west side, 
the inside. 

the wood at the outside, 
it is uppermost. 

^Anthropos U (1931) 485; Philip. Jour. Sci. 71 (1940) 21. 


The Philippine Journal of Science 


IV. Many adverbs of place are formed by substantives in the 
oblique and by the preposition ididy followed by a substantive, 
Examples : 

napdnda iti rudr 
napdn iti adayo 
awdnda iti uneg 
adda iti malikuddk 
addd iti masangoanak 
adddda ididy sirok 
inka ididy rudr 
immuli ididy ngato 
makaddnon ididy babd 
nagydn ididy tengngd 
inkayo ididy uneg 
naidisso ididy rabdwna 

they went outside. 

he went far. 

they are not inside. 

it is behind me. 

it is before me. 

they are below. 

go outside. 

he climbed up; he ascended. 

he gets down. 

he stayed in the middle. 

go inside. 

it was placed on top. 


I. 1. There are two adverbs of negation : sadn and di. 
a. Sadn is connected with the word it modifies by the liga- 
ture a or nga. Examples: 

saan nga aso 

sadn a ni Pidro 

sadn a nasaydat 

sadn a nalagdd a baldy daytd 

sadn nga dmvn 

sadnda a tallo 

sadn a daytoy 

sadn a napdn 

sadnka nga immdy 

sadnda a kaydt 

sadnkanto a di mapdyat 

it is not a dog. 
not Peter, 
it is not good, 
that is not a strong house, 
not all. 

they are not three, 
not this, 
he did not go. 
you did not come, 
they do not like. 
I shall certainly tread it 


b. Di precedes it immediately without any ligature. The pe- 
culiarities connected with the use of the possessives, when di is 
followed by a substantive or a substantival verb, have been 
alluded to before. Cfr. The Pronoun, II. Possessives. 

2. Peculiarities. I.^^ Examples: 

dida gayySm 

diak mapdn 

dikam kaydt 

dikay umdy 

didakami ayaten 

dina pay nalpds ti nagsao 

he is not their friend. 

I do not go. 

we do not like it. 

don^t go. 

they do not love us. 

he had not yet finished speaking. 

•Anthropos28 (1933) 691-692. 

Vanoverbergh: lloko Adverbs and Prepositions 181 

3. Saan is the most extensively used. 

Di is rarely used by itself without an accompanying pronoun, 
except in some expressions and in literature, but it is generally 
preferred to sadn when the verb is in the imperative. Ex- 
amples : 

dipay la umciy doesn't he come yet? 

di naimbdg daytd is that not good? 

asino ti di agaydt who won't be glad? 

sumurot kenkudna a di agtantdn he follows him without delay. 

dika mamapdtay thou shalt not kill. 

dika agtdkaw thou shalt not steal. 

Note 16. Ammok pay and ammok kadi or ammo pay and ammo kadi 
are strange expressions meaning "I don't know, it cannot be known". 

Note 17. Naymdn, "who knows?", has become antiquated. 
^ II. "Never'* is translated by adding the adverbial expressions 
uray kaano (man) or uray and (man) to a negative sentence. 
After uray may be placed: no, inton (regular forms) adton, 
atton (rarely used). Examples: 
diakto gatangen uray kaano 

^«w I shall never buy it. 

sadnna a kaydt uray no and man he never wants it. 
dikamto gayyem uray no and he will never be our friend. 
uray inUn kaano man never. 

Uray and sadnak pay a nangdn I never ate bread. 
iti tindpay 


I. In general, an interrogation, in lloko, is distinguished from 
a statement merely by the inflection of the voice. 

II. Besides several adverbs of interrogation which have been 
studied under the Number, the following are the most used 
interrogatives introducing a sentence in lloko. 

1. Kaano ti (ordinary form) and iti anid ti (rarely used), 
"when T* Examples : 

kaano ti isasangpitna when did he arrive? 

kaano ti yadymo manin when will you come again? 

kaano ti panagsardengda when did they stop? 

kaano ti pannanganyo ditoy when do you eat here? 

kaano nga alddw ti panang- on what day will you bring it? 


iti anid nga alddw ti ipapdnna on what day will he go? 

2. Intonano ti (most common form), inton kaano ti, tonano ti, 
ton kaano ti, attonano ti, adton kaano ti or atton kanno ti, 
antonano ti, anton kaano ti, "when?" (future). Examples: 

intonano ti yadymo when will you come? 

intonano ti kaydtyo a pannaka- when do you want it made? 


The Philippine Journal of Science 


intonano ti kaaddd ti ndkemda 
inton kaano ti panangyegda 
tonano ti pannakasdpulna 
ton kaaim ti pannagndda 
attonano ti kondyo 
atton kaano ti panagkaydtda 
antonano ti panagdpit 
anton kaano ti panagbdyadmo 

when will they have sense? 
when will they bring it? 
when is it needed? 
when will they walk? 
when do you think? 
when will they like it? 
at what time is the harvest? 
when will you pay? 

Note 18. Nowadays these forms are very often superseded by kaanonto. 

kaanonto ti panangiserrekna 
kaanonto ti panagodongmi 

when will he bring it in? 
when will we go to town? 

3. Adin (o) ti, sadin (o) ti, "when?'' Examples: 

sadin ti papandnTia 

sadino ti nakabirokam kada- 

adin ti nagtaraydn ti dso 
adino ti paglutodnyo 
sadin ti kaydtda a pakaikabi" 


where does he go? 

where did you find them? 

where did the dog run? 
where do you cook? 

where do they want it located? 

4. Asin (o) ti, si asino ti, "who?'* Examples: 

asino ti maydt 

asin ti piUSm 

asin ti lakdy nga immdy idi kal- 

si asino ti napdn 
si asinoak tapno daydwendak 

who is willing? 

whom do you choose? 

who was the old man who came 

who went? 
who am I that you should honor 

5. Ania ti (ordinary form) and ti, 'Vhat?'' Examples: 

anid ti kondm 
anid ti kaydtmo 
anid ti pagsdgadko 
anid ti gindtangda 
anid ti ildbasyo 

what do you say? 
what do you want? 
with what can I sweep it? 
what did they buy? 
with what do you pass? 

6. Ania ti gapona, dpay (ordinary forms), apay-dpay (mostly 
used in literature), apayd (rarely used), "why?'' Examples: 

ania ti gapona nga immdyka 
anidt gapona a madida 
anidt gapona nga awdn ti maydt 
anid ti gapona a nabartek 

anidt gapona a bimmassit ti 

dpay a sukd ti inkdbilyo 

why did you come? 
why do they refuse? 
why is nobody willing? 
why is he drunk again? 

why did the wine diminish? 

why did you put vinegar? 

Vanoverbergh: Iloko Adverbs and Prepositions 183 

dpay a didakami kitkitden why don't you look at us? 

dpay nga dso ti ginatangd't why did they buy dogs? 

dpay nga awdn ti sumangpet why does nobody arrive? 

dpay a sikd ti baoninda why do they send you? 

apaydpay mapdnkayo why do you go? 

apayd umdy ^hy ^^es he come? 

1. Kasano ti, an-anoen, agm-ano (past: nagan-arw), "how?" 

Examples : 

kasand ti pannakaardmidmt how was it made? 

kasano ti panagtardyko no nasa- how can I run if my foot hurts. 

kit ti sdkak -, , , j; 4.i,„„9 

kasand ti panangbirokko kada- how can I look for them? 

kudda _. j„Lj.- :f 

an-ano6n a bayddan ti utang no how can one pay ones debts if 

with that? 
agan-andkam a bumaknang how can we get rich? 

nagan-anokaarimudr how did you get out? 

8. AgpmrM (past: nagpamo), "in what direction?" Ex- 
amples : 

agpaano daytd lakdy in what direction does that old 

man go? 
nngpa^trwkayo idi kalmun in what direction did you go yes- 


9. Tagmno (ordinary form). yan0, "from where?" Exam- 

pies : 

, - .,..^. from where are those ones? 

. 'j.^..^ from where are your 

tagawmkayo ^^^^^ ^^^ y^„, 


NOTE 19. The same question may be asked this way: Anid ti ilim? 
"what is your town; from where are you?' 

III. Most Of these interrogatives may be used either ^«;'?- 
verbs of interrogation or as conjunctions nitroducmg an mci- 
dent^l clause, in which latter case they are preceded by the 
simple conjunction no. Examples: 

amm^k pay no kaan6 ti yadyna I don^t know when he camj 
ibagdm no intonano ti panang- tell me when you wui s 

dZZ^^tdin. ti napa- I ask^ where he went. 
saludsMem no asino ti mayat ask who is willing. 


The Philippine Jommal of Science 


ammoda no si asino ti nagtdkaw 

isurona no anid ti ddlan 

ipalawdgna no anid ti gapona a 

inpaddmagna no dpay nga awdn 
ti immdy 

dikayonto panunoten no kasano 
wenno anid ti saoenyonto 

kitdenyo dagiti ubbing no ka- 
sano ti idadakkelda 

they know who stole. 

he shows them where the road is. 

he explains why they refuse. 

he told me why nobody came. 

take no thought how or what to 

look at the children how they grow. 

Note 20. Formerly the adverb nga was generally used to introduce an 
interrogative sentence, but actually it has become entirely obsolete, and 
it only exists in the complex adverb ngdman. 

IV. ffgay, aya and ngamin are adverbs of interrogation which 
nearly always follow some other word. fTgaman or ndman, 
which is not used very extensively, stands by itself or introduces 
the sentence. 

1. Ngay is a simple interrogative. Examples: 

why do you refuse, what is the 

why, what is the matter? 

what are you looking for? 

where were you yesterday? 

what right has he? 

where did he go? 

and John? (where is he; where is 
his share?) 

and your studies? (when talking 
to a boy or a girl who wants to 
marry, for instance) 

why shouldn't there be any? (an- 
swering contradictorily a person 
who said there wasn't any) 

Note 21. The form ay for ngay has become antiquated. 

2. Aya and ngamin are interrogatives implying an additional 

dpay a madika ngay 

dpay ngay 

anid ti birokem ngay 

sadin ti ydnmo ngay idi kalmdn 

anid ti kalintegdnna 'ngay 

napandnna ngay 

ni Judn ngay 

ket ti panagddalmo ngay 

sa pay awan ngay 

notion of wonder. Examples: 

napdnak ayd 

sidk ayd ti baonenyo 

inka ayd 

pudno komd ayd 

aldem man ayd 
anid ketdin ayd 
nalukmeg ayd 
dpay sakit ayd ngamin 

did I go? (implying the contrary) 
is it I you send? (I thought 

somebody else) 
do you go? (I thought you 

wouldn't or not yet) 
it should be true, wouldn't it be 

all right, take it (why not?) 
stop it (or: what can be done?) 
h he really fat? 
why, is it sickness indeed? 

Vanoverbergh: Iloko Adverbs and Prepositions 185 


sidk ayd ngamin 

sadnda ngamin a sangap'alo, da- 

giti nadalusdn 
addd ngamin nagkurang kada- 

bulsekkami met ngamin 

mabalinna ngamin ti agsubli 
ngamin komd no addd dsok 
mabalinda ngamin ti agliddy 
dakdakkelda ngamin ngem 

iti 'ngamin Abra ti paggapodn ti 

namdti ngamin kenkudna ti 

maysd kadakudda 
sadn ngamin a daytdy ti andk ti 

itedmo ngamin ti kankansn 

kastdy ti nagtutuldgantay nga- 
dgpayso ngamin 
awdn met ngamin pirdk 

apayd ngamin a ditay ipatpateg 
ti kukuatayo 

I? {do you suppose I did it, not 

at all) 
were not ten made clean? (and 

why do I aee only one?) 
were you really in need of some- 
are we blind indeed? (do you 

consider us as blind?) 
can he go back? (I think not) 
I wish I had a dog. 
can they grieve? (I think not) 
why, indeed, they are taller than 

my father, 
the timber comes from Abra (who 

would have thought that?) 
one of them believed in him, isn't 

it strange? 
was not that the carpenter's son? 

you are giving sweets (we didn't 
know that) 

did we agree to such a course of 
action? (I thought the contrary) 

is it true, really? 

well, there is no money (we 
thought there was) 

why indeed don't we prize our be- 

3. iTgdmdn or ndman means "why?'* and so on. 


ngdman apo nga awdn ti nd- 

kemko tapno isn ti aramidek 
ngdman itd a di met naipdkat 

itoy dagdtayo 
ngdman pay nga ikkdnnak nga- 

td ti pagduaduadk kenkuana 
ngdman pay a marikndk daytd 
ngdmansa no gumdtang ditoy 

ti pdgay 

Examples : 

what is the matter? why? 

why, Sir, do you think I have no 

sense that I should do that? 
why is it now that it was not given 

to this country of ours? 
why do you seem to make me 

doubt him? 
why do I feel that? 
what if I buy rice here? 

Note 22. The interrogative ma«, *Vhy?" which follows the word it 
modifies, is very little used nowadays, except in the expression sadn 
manf "why not?,*' "certainly," when answering affirmatively a negative 
question or its equivalent, and in the complex adverb ngdman or namdn. 
Examples : 

dika man mamdti why don't you believe? 

sadnka a wapdn ayd? — Sadn didn't you go really? — ^I went 

man though. 

mdnkansa a napdn? — Sadn man you didn't go I believe? — ^I went 


186 The Philippine Journal of Science 


1, To indicate continual repetition of an action, the word 
expressing the action is twice repeated and the abverb la from 
Ideng followed by the ligature a or nga connects the two. 
Examples : 

sarita la a sarita daytd lakdy that old man does not stop talking. 

saoka la a sao you are talking all the time, 

humassit la a bumassit ti bagds the rice becomes less and less. 

lumukmeg la a lumukmeg ti the child grows fatter and fatter. 


umadH la nga umadii ti dodon the locusts increase in number all 

the time. 

sdngitda la a sdngit they are always weeping. 

jmmMawda la a pumMaw they become whiter and whiter. 

ngumisit la a ngumisit ti rttpam your face becomes blacker every 


tardy la a tardy running all the time. 

lumdlo met a lumdlo it becomes worse and worse. 

sidk la a sidk ti baonenyo, you always send me. 

II. The concept of completeness, purity, and so on, is expressed 
by the following words, which are connected with the word 
they modify by the ligature a or nga: Pasig, biig, bin4g (pre- 
ceding) and puhs (following or preceding). Examples: 

pdsig a tuldng all bones. 

biig nga allid pure wax. 

bin-ig a balitok pure gold. 

puUs a sinamdy all of it sinamay. 

awdn a pulos there is nothing at all. 

III. 1. "So" is rendered into Iloko by the indefinite adjectives 
in kas, kastoy, kastd, and so on. Examples: 

aramidem a kastd do it so. 

kastoy latta like this all the time. 

kasdidy a natdyag as tall as that; so tall. 

kastdy nga adtl as much as that; so much. 

kasdi a bassit as little as that; so little. 

kastoy a dakkel as great as this; so great. 

2. "Equally" is expressed by pdda or agpdda (past: nag- 
pdda) with ligature a or ng^a following. Examples. 

ti pddana a natangsit his equal in pride. 

nagpadpddada a nadagsin they were equally heavy. 

agpadpddada a nalag-dn they are equally light. 

dagiti pddami a napaiTgldw our equals in poverty. 

Note 23. The latter is an expression the Iloko generally use, instead of 
saying "the poor" and so on, whenever they refer to a quality which is 
humiliating to their neighbor. It would be rather bad taste to say that 
a man is poor, without referring to him as one's equal in poverty, even 
though one be as rich as Croesus. 

dagiti pddami a nanengneng our equals in stupidity. 

Vanoverbergh: Iloko Adverbs and Prepositions 187 

IV. Other adverbs of quality and manner are: 

1. tlnay, **very much" (following, no ligature)! Examples: 

dakkel unay it is very great. 

bassit unay very little. 

kaydtko unay I like it very much. 

ipapilitna unay he insists much. 

magna iti nadards linay he walks very fast. 

Note 24. Bassit means "a little" and so on. Examples: 

dakkel bassit rather great. 

adu bassit rather much. 

nadagsdn bassit rather heavy. 

adayo bassit a little far; rather far. 

2. Amangdn (ordinary form), amamangan, allek (rarely 
used), "how great'' (preceding). Examples: 

amangdn a nag adu ti tdo how many men. 

amangdn a nagrikiit unay how very difficult it is. 

amangan a salditmo ngatd a looking at you I think you must be 

buybuydek exceedingly well-favored. 

amangdn imbdg ti baldyyo how good is your house. 

3. (N^an^) nan^roTMi, sarigrfcanj^rona, ''especially." Examples: 

daytoy ti nangrona a ddkes this is especially bad. 

nangrona Unay iti bigdt especially in the morning. 

isu ti nangnangrona a bassit he is especially small. 

sangkangronada a napintds they excel in beauty. 

Nt)TE 25. This adverb has a form of relative superlative: kangrondan. 
Examples : 

ti kangrondan a tuleng the deafest of all. 

ti kangrondanda a ddkes the worst of them all. 

4. Istdy, nagistdy, nagistaydn, "nearly" (preceding, no lig- 
ature). Examples: 

istdy natndg he nearly fell down. 

nagistdy naitibkM he nearly stambled. 

nagistdyak natdy I nearly died. 

nagistaydn pimmdnaw he was nearly gone. 

5. iTganngdni di, nganngdni sadn, apdman (ordinary forms), 
apagapdman, apaganamd, "scarcely" and so on. Examples: 

nganngdni di agiini it scarcely sounds. 

nganngdni sadn a mangngeg it is scarcely audible. 

apdman a nangted he barely gave. 

apdman a nagsdgadda they hardly swept it. 

apdman addd there is just a little. 

apdman ta maydt he is but just willing. 


The Philippine Journal of Science 


6. No maminsdn, *in a way'' and so on (no ligature). Ex- 
amples : 

no maminsdn nasaydat 
no Tnaminsdn kaydtko 
napintds no maminsdn 

it is good in a way. 
I more or less like it. 
it is beautiful in a way. 

7, Sigiid, 
amples : 

'originally'' and so on ligature a or nga. Ex- 

sigud a ddkes 
sigud a bulsek 
nabakndng a sigud 
sigud nga awdn ti kukuana 
sigud a nakas^rat 

it is intrinsically bad. 

he was bom blind. 

he has always been rich. 

he never had any property. 

he had always been able to write. 

8. I0ray {no) kasano, uray (no) ipapaano, "anyhow, anyway" 
(no ligature). Cfr. VIII. Other Adverbs. I. 3. 

9. Mahay dg (past: nabaydg), kalkald (pay) (ordinary 
forms) f makamano, makamanmano (rarely used), ** a long time" 
and so on. Cfr. II. Adverbs of Time. III. Note 9. Examples : 

mabaydg ti panangpdtitda 
addd a nahaydgen 
nabaydgak a nangdUm 
kalkald pay ti isasangpitko 
kalkald pay ket dika pay nalpds 

they ring the bells for a long time, 
he is here a long time already. 
I took some a long time ago. 
I arrived a long time ago. 
sucn a long time and you have not 
yet finished it. 

10. Apagbalikds, apagdarikmdt apagkanitd, apagkigmdt, 
apagdipds, sadn a kakigmdtan, "in a short time" and so on. 
Examples : 

mdluto Hi apagbalikds 

rimmudr iti apagdarikmdt 
sumangpetto iti apagkanito 
nakarubbudt iti apagkigmdt 

apagdipds a miibus ti sabd 

sa^n a kakigmdtan adddn 

11. Pagamodn, 
ture) . Examples : 

it can be cooked in a very short 
time (the time of pronouncing 
a word) 

he came out in a moment. 

he will be here in a moment. 

he was ready in the twinkling of 
an eye. 

the bananas were consumed in a 
very short time. 

one moment and here he is al- 
ready (before the eye twinkles) 

*at once" and so on (preceding, no liga- 

pagamodn addd 
pagamodn nagtdlawen 

it appears at once, 
he was gone unawares. 

Vanoverbergh: Iloko Adverbs and Prepositions Igg 

12. Dards "quickly" and so on (following, no ligature), 
Examples : 

maibiis dards 
agsubli dards 
umiilogkat dards 
makalipat dards 
aramidem dards 

it is used quickly, 
he returns quickly, 
go down quickly, 
he forgets easily, 
do it quickly. 

13. Manen, "again''. This adverb either stands by itself or 
follows immediately some other word. En may be separated 
from man and joined to some other word further down in the 
sentence, and then it follows the rules of the simple adverb 
en. Cfr, II. Adverbs of Time. IV. 

II. Adverbs of Time. IV. Examples: 

napdn manin 

dpay a nabartekka manen 

sikd Tnanin 

addd man daytd lakdyen 

nagsakit man idi kalmdnen 

kasdi wMn met Idengen 

napdn man Tuigtdkawen 

adddtay man dminen 

awdn man ti makitamon 

nagsubli man kanidkon 

addd man iti bidngmin 

he went again, 
why are you drunk again? 
you again? 

that old man is there again, 
he got sick again yesterday, 
just like that again, 
he went again to steal, 
we are here again all of us. 
don't you see anything again? 
he came back again to me, 
he is again under our care. 

14. Pill, with the meaning of "what should not be", is rarely 
used (preceding, no ligature). Examples: 

pill ket dida sapUten ti nasubeg 

pili natdy ti bakndng ditoy ili- 

pili naboon^da ti kapintdsan a 



why don't they rather whip the 

very disobedient one? 
the rich man of our town died, 

why he? 
they must break just the nicest 



I. The concept of excess is expressed in Iloko by; 
1. The term napdlalo or nalabes. Examples : 

napaldlo a dakkil 
napalaU a nasadut 
ddkes a napaldlo 
imbannogka a napaldlo 
umintim iti napaldlo 
mangdnda iti napalpaldlo 
nalabes ti nginana 

too big. 

too lazy. 

too bad. 

you are too tired. 

he drinks too much. 

they eat too much. 

it is too expensive. 


The Philippine Journal of Science 


he is too good a man to think 

about revenge. 
I love you too much to be able to 

look at you without embracing 


2. The adverb nnay following immediately the word it mo- 
difies and the conjunction tdpno introducing an incidental 
clause. Examples : 

naimbdg iinay a tm tapno mapa- 

nunotna a balsen 
ay-ayatenka unay tapno maha- 

linka a buybuyden a di araku^ 


II. "Only'' is expressed in Iloko by: 

1. The adverb Ideng, which always immediately follows some 
preceding word without connection by ligature. When another 
word follows, it is often abbreviated into la. Examples: 

sidk Ideng I only. 

ilray ti Ideng panangsdgidko even only my touching him. 

isMa Ideng ti pangikkdnna he gives only to them. 

daytoy a ragsdk sa Ming marag- this joy will only be obtained after 

pdt iti kalpasdn ti adu a pa- 

tdo la ti addd 
agbidang latta 

much sorrowing. 

only men are there, 
she just wears a tapis and that's 
sdngit la ti ammona he knows only weeping. 

Note 26. This adverb is extensively used in Iloko. We have seen some 
instances before and we shall see more of them in due time. Sometimes 
the change it effects in the meaning of the sentence is very slight and 
often practically nil; therefore we shall give some examples of its use 
here. Examples: 

sidk Idengen 
awdn la ti umdy 
matHrogka Ideng 
Urayka Meng 
napudot pay Ideng 
makitam la no addd ididy 

di pay la pimmdnaw iti baldyna 
kondk Ian nalaklakdda 

diak la idd napagraSm 

masdpulmi la iinay 
inka Ideng 
addd pay Ideng 

awdn pay Ideng 
adH pay Ideng 

let me do it. 

nobody has come yet. 

go on sleeping. 

it depends on you. 

it is still warm. 

you will see it if he is there (that 

it is worse than what you 

he has not left his house yet. 
I thought though that they were 

I was wholly unable to have them 

respect him. 
we need it very badly, 
all right, you can go. 
he is still here; there is still some 

he is not here yet. 
there is still much left. 

2. The reduplication of the initial, open syllable of the stem 
with the next consonant, if there is any, of the personal pronouns 

Vanoverbergh: Iloko Adverbs and Prepositions 191 

of the first series, of the complete forms of the demonstratives, 
of some numbers (cardinal numbers, distributives, and so on), 
of the prefix sariga and so on. Examples : 

sisiak I only. 


we only. 


only this one. 


only these ones. 


four only. 


only eight. 


he is alone. 


they have only five each. 


only ten each. 


only one jarful. 

dMua ideng a ddlan 

only two roads. 

banbannogda ti mamagbagd 

it is useless for them to give 

advice, (literally: only their 

labor in giving advice.) 

Note 27. The forms datdaytoyj datdaytd, and so on, instead of daydaytoy, 
daydaytd, and so on, sometimes occur. 

3. The reduplication of the first or of the second part of the 
complex prefix of the multiplicatives ; in the latter case mami 
and mamin become maminpi and maminpin respectively. 
Examples : 

namnaminsdn only once. 

namnaminpulo only ten times. 

maminpinsdn only once. 

maminpitlo only three times. 

maminpinsidm only nine times. 

Note. 28. Maminpisdn may also mean "completely". Example: 
pariitekto a maminpinsdn I shall grub it up completely. 

III. "A little'' is rendered into Iloko by: 

1. The term bassit following the word immediately without 
any ligature. Examples: 

natured bassit he is rather bold. 

addd bassit there are just a few. 

mangdn bassit he eats a little. 

yadayona bassit iti dagd he puts it a little distance from 

the earth. 

2. The term addd or awdn followed by the principal word 
in which the initial, open syllable is reduplicated together with 
the next consonant if there is any. Examples: 

addd tultHlengna he is a little deaf. 

awdn ti taktakrotda they are not timid at all. 

8104 5 


The Philippine Journal of Science 


addd pangpangkismo 
addd samsam-itna 

you have a cast in the eye. 
it has a tinge of sweetness. 

Note 29. The prefix kanika adds the meaning of "less than" to a 
number. Cfr. The Number, I. Cardinal Numbers. IV. Note 1.^^ Ex- 
ample : 

kanikapdt less than four, 

IV. The multiplicatives and the distributives, which were 
studied under the numbers, are connected with the word they 
modify : 

1. By the ligature a or nga, when they precede; 

2. By the article in the oblique, when they follow. Examples : 

namindud a napdn 
maminpdtkami a mangdn 
fmmitloda a nagaydh 
naminwaloda nga agtardy 
saggaysdda nga agsubli 
sagpapdtda a napdn 
nagtardyak iti nawinsdn 
gumdtangkaminto iti mamitlo 
nagikkisda iti namindud 
agulbodda iti maminribu 
nagsdngit iti kapamitUna 
iiangdlada iti sagdudud 

he went twice. 

we eat four times. 

they called three times. 

they run eight times. 

they come back one by one. 

they went four by four. 

I ran once. 

we shall buy three times. 

they shouted twice. 

they lie a thousand times. 

he wept for the third time. 

they took each two. 

1. Adverbs implying affirmation, confirmation, and so on: 
Examples : 

1. Wen, "yes*' (used by itself), 

2. Pay, "yet, also" and so on (following, no ligature). Ex- 
amples : 

ad4 pay 

addd pay 

agsao pay Ideng 

ni pay Ana 

nauyong pay 

dakkil pay 

bumulodak pay 

ikkdnnak pay 

bagds pay 

sidk pay 

diak pay nangdn 

sadn pay a napdn 

awdn pay 

di pay dimting 

mangdnka pay sdka mapdn 

di pay nasaydat 

there is much left. 

there is some more left. 

he still talks. 

Ann also. 

he is also severe. 

it is also big. 

I want to borrow more. 

give me some more. 

rice also. 

I also. 

I didn't eat yet. 

he did not go yet. 

he is not here yet. 

he didn't arrive yet. 

eat first and then go, 

is that not good? it is really good. 

^^Anthropos 28 (1933) 711-712. 

Vanoverbergh: Iloko Adverbs and Prepositions 193 

nakigtotak pay 


pay no amnvok a 

I was startled (pay corroborates 

the statement), 
would I do it if I knew it were 


3. Agpapdn pay, uray, ''even, whatever" and so on (preceding, 
no ligature). Cfr. Conjunction "Although'\38 Examples: 

agpapdn pay sangalubongan 
diak kinita agpapdn pay ti riipa 

ni amdm 
uray isu napdn met 
Uray dso awdn 
uray sidk 

awdn natdy nga Uray maysd 
Uray no kaano ti kondm 
nasaydat ijiray kaxino 
umdykayo uray no and 
uray dagitd dsoda met 
Uray and diak kaydt 
uray no sadin ti ydnda mabiro- 

addd iti Uray sadino a yan ti 

uray no si asino 
ayatin ti Uray si asino a maka- 

uray si asino ti addd ayabdm 
iHray no anid ti kondna 
Uray anidt kita a tagildko 
uray anid a sakit nga addd ken- 

madiak uray no kasano 
iLiray ipapaano ti umiggim 
uray kasano ti panangay-dyona 


even the whole world. 

I did not see even your father's 

even he, he went also, 
there is nothing, not even a dog. 
I also. 

nobody died, not even one. 
whenever you say so. 
it is always good, 
come any time. 

even those dogs of them, they also. 
I don't like it at all. 
wherever they are we shall find 

it is all over the world. 


whoever sees it loves it. 

call whomsoever is there, 
whatever he says, 
any kind of merchandise, 
whatever is his ailment. 

I won't anyhow, 

howsoever we can take hold of it. 

howsoever he coaxes me. 

4. Mgarud: confirmatory (ordinarily following, no ligature). 
Examples : 

inka ngariid 
mapdnak ngaHiden 
inbagdk ngarHden 

ti ngariid kinondna kastoy 

go now (your reasons convinced 

me; do as you told me). 
I am going now (as I told you; 

as you told me). 
I said it (why do you say I did 

not; why didn't you believe me; 

I told you so, as it had to be 

now, what he said was like this. 

» Unpublished. 


The Philippine Journal of Science 


sidk ngaruden ket addd pay 

umdy ngarud 

anid ngarud it kayatyo 

awdn ngarud 

sikd ngaruden 

ni Juan ngarud ti ayabdm 

uminandka ngarud 
puorak ngarud daytd kayo 
sadnyo ngarud a kaydt 

dpay ngarud a makitam 

into ngarud no umdy matari- 

mdanto dmin 
ti ngarud lakdy ti nangardmid 

it is I, and still you have some- 
thing to say. 
he is here now (stop your mouth) ; 

of course he comes, 
what do you want (why don't you 

of course he is not there; he is 

not there indeed, 
now you (I can't; you do it as 

you pretend that you can), 
then call John (if you cannot find 

another one), 
take a rest now (as you are tired), 
if so I shall burn that wood, 
why, you don't like it (why do 

you object to my taking it?) 
why, do you see it? (you said you 

were blind), 
consequently, when he comes, 

everything will be settled, 
of course the old man made it; it 

was the old man who made it 

(nobody else), 
then what will happen tomorrow? 
whosoever heard it was astonished 

(it couldn't be otherwise), 
all right (stop asking me for it); 
yes, isn't it strange? 

5. Sgad: abbreviation of the preceding, mostly used in liter- 
ature (following), no ligature. Examples: 

anid ngarud ti adddnto no bigdt 
dmin ngarud a dimngeg nasdd- 

wen ngardd 

ta nadalUs ngad dmin 

anid ngad ti gapona 
dika ngad tuldden 
ti ngad agturdy ti rehhengtay 
nga annugoten 

because everything was clean of 

then why? 

thus, don't imitate him. 
thus, we have to obey rulers (no 


6. Piman: confirmatory^ ordinarily implying commiseration 
(ordinarily following, no ligature). Examples: 

ta awdn pina^n ti nakapanunot 

andknaka piman ti okom ket 

isM gapona a kastd ti ard- 

ta babbdi ket piman dagiti ub- 

ta is'A piman awdn pay inar- 


as there was nobody who thought 
(it's too bad). 

because you are the son of the 
judge, is that a reason to be- 
have as you do? 

as the poor children were girls 
(what could they do?) 

because he hadn't done anything 
yet, poor man. 

Vanoverbergh: Iloko Adverbs and Prepositions 195 

vmduadud piman no addd ken 
panagtag taguinep 

piman ni Maria 

kdsnak la andk piman 

dina kaydt ti mangpabain piman 

mabain metten piman a mangH- 

ti andkna a siisemy piman siisem 

ta dina pay la ammo ti napa- 

sdmak ti amdna 

he is really in doubt if he is 
dreaming or not (the news is 
so very bad). 

poor Mary. 

she treated me as a son, poor girl. 

he doesn't want to make her feel 

he is ashamed now to repeat it, 
poor boy. 

his son who was smiling, smiling 
indeed, as he did not yet know 
what happened to his father. 

7. Da: confirmatory (no ligature). Examples: 

da ikkdnnak da 

innak da 

da man dinak kikien 

now, give me some, quick. 

I go now. 

leave me, don't tickle me. 

8. A: confirmatory (at the end of a sentence). Examples: 

inkayo a 

mangtedka a 
isu a 

awdn a 

modi a 

makaammoka a 

addd a 

kastdn a 

wen a 

adu a 

birokem a 

isuda ti pagdamdgam a 

no mapdnda dmin Uray sidk ma- 
pdnakto met a 

go now (why do you wait, I don^t 

keep you back), 
give me some now, won't you? 
that is he of course; all right then, 

if you say so. 
there isn't any of course, 
of course he won't, 
that is your business of course, 
he is there of course, 
so long, 
yes, of course, 
there is much of course, 
well, look for it. 
ask them (why do you bother 

if they all go, I also shall go, 

shouldn't I? 

II. Adverbs implying opposition: 

1. Ket, ketdi, "rather'' and so on (following, no ligature) 
Examples : 

addd ket 

sabdli ket ti rnalagipko 

isut sadn ket a sabdli 

awdn pay met ket ti nasasaona 

kas ket bassit ti pategna 
ngem kinondna ket kenkudna 

there is some (why do you say 

there isn't any?) 
now I am remembering something 

he it is and not somebody else, 
he had not said anything yet 

as if it were of little value, 
but he said to him though. 


The Philippine Journal of Science 


a kas ket sakiten ti ndkemda 

ngem kanenda ket ti tindpay 
inardmidda ket kenkudna ddgup 

ti kinaydtda 
nadlana a di ket naslep 

dina ket maibbatdn 

awdn ket a pulos ti linabdsanna 

mvdn ket kenkudna ti uray anid 

a ddkes 
awdn ketdi 

sikd ketdi 
nagdaksangdsatak ketdin 

kondm ketdi apoy makitam la no 
addd ididy 

wen ketdi ngem nanginansa 

dida la ketdi mabain 

wen ketdi no maydt ti bakitko 

isunsa ketdi daytoy ti birbirO' 

adddnsa ketdi nangyeg ken- 
kudna iti pirdk 

kaydtyonsa met ketdi ti agbalin 
a pangolo 

they seem to be sorry about it 

but they eat the bread though. 

but they treated him howsoever 
they liked. 

he took it without getting wet at 

he doesn't let it go though. 

he didn't skip anything at all 

there isn't anything bad in him 

there isn't any (why do you de- 
ceive me; why don't you believe 

you better do it. 

what bad luck is mine (complain- 

that is what you think, sir, but if 
he is there, you'll see. 

all right, but isn't it expensive? 

why, they should be ashamed. 

all right, if my wife agrees, 

isn't this what you are looking 

hasn't somebody given him money? 

don't you act as if you wanted to 
become chiefs? 

Note 30. Get has become antiquated. 

2. Met, met Ideng, met la "also, though'' (following, no liga- 
ture). Examples: 

my brother is here also (he might 

not have been, but he is), 
but he is not there (didn't you 

say he was?) 
he is there though, 
he refuses though, 
now he does it at once, 
it is not very far though. 
I also (don't leave me out), 
also the dogs (don't exclude them), 
well, there is no money (we 

thought there was). 
Lewis also, 
we are blind indeed (do you think 

my wife isn't here yet (I thought 

she was). 

addd met ni kabsdtko 

awdn met 

addd met 

madi met 

dagtisenna mettin 

di unay adayo met 

sidk met 

dagiti dso pay met 

awdn met ngamin pirdk 

ni met Luis 
bulsekkami met ngamin 

ti asdwak auin pay met 

Vanoverbergh: Iloko Adverbs and Prepositions 197 

simmangpet metten 

asid4g met 

sikd pay met 

awdn pay met ti m^zydt 

he has arrived already (don't 
think he hasn't). 

it isn't so far. 

why, you also? 

nobody is willing yet (although 
you said otherwise; although the 
salary is very high). 

nobody is willing though. 

he isn't there though. 

he is here though. 

he refuses though. 

he did his best though to get rid 
of it (sorrow, and so on) quick. 

just as formerly. 

he is in that same place. 

it is the same. 

that same thing. 

what we call pugot (black one) 
is just a man. 

they were able to see on that 
exact moment. 

Note 31. "Self," as applied to persons, may be expressed either by the 
adverb met laeng or by the term hagi or hagi met Ideng. Examples: 

awdn met la ti maydt 

awdn met Ideng 

addd met Ideng 

madi met Ideng 

inagawdanna met la nga inliw* 

liwag dards 
kas met la idi 
daydi met Ideng ti ydnna 
tsti m,et Iden^ 
daytd met Ideng 
ti pugot isti met la ti tdo 

idi met la kanito nakakitada 

ti bagina komd ti panunotenna 
pinatdyna ti bagina met Ideng 
ayatentayo ti bagitayo met Id- 
sidk met Ideng 
ni dma met Ideng 
ni met la Ana ti napdn 
isuda met la ti agbHsor 

he should think of himself, 
he committed suicide, 
let us love ourselves. 

I myself. 

my father himself. 

Ann herself went. 

they themselves are enemies. 

III. Adverbs implying doubt: 

1. iS^ata, "perhaps'' (following, no ligature). Examples: 

addd ngatd ni apo lakdy 

awdn ngatd 
sikd ngatd ti nangdla 
umdyakto ngdtd 
nayad-adddna ngatd ti nangdn 
sidk la ngatdn ken sikd ti adddn 

a sipup&ot 
agan-anoak ngatd no sumangpit 

no kastd komd amangdn ngatd, 

ti takning dagiti agturdy 
amangdn a gagetmo ngatd ti 


is the old gentleman there per- 

maybe he is not there. 

did you take it perchance? 

perhaps I'll come. 

maybe he only came to eat. 

maybe you and I were the only 
ones who noticed it at the time. 

what can I possibly do if he ar- 
rives here? 

if it were so how virtuous the 
rulers would be. 

isn't the diligence I notice in yo» 
very great? 


The Philippine Journal of Science 


ididy baldy iigatd ti ydnna 
nalukmSg ngatd la Unay itdn 
ammo ngatd ti kabsdtko 

maybe he is in the house, 
maybe he is very fat by this time, 
maybe my brother knows. 

2. Sa, "probably'' (always joined to a pi-eceding word; the 
ligature n connects the two if the preceding word ends a vowel). 
Examples : 

adddnsa apo 

adddnsa pay anndkda 


kaydtnansa ti sumrik 


ninsa Pidro (ni Pedronsa) 

isunsa ti nagtdkaw kadagiti hd- 

uppdtsa ti nabdti 

I think he is there, sir. 

haven't they also children? 

I don't think there is any. 

he probably wants to enter. 

I think so. 

Peter probably, 

I think that is the one who stole 

our pigs. 
I think there are four left, 
they are asleep probably. 

3. NoMMt, 
Examples : 

'probably" (preceding, ligature a or nga). 

naMbit a simmangpeten 
naMbit nga awdn 
naldbit a kastd 
naldbit nga ayatSnnaka 
naldbit nga addd pirdkman 
naldbit a due puloda mettSn 

he probably arrived already, 
it is profable that there isn't any. 
that is very likely, 
he probably loves you. 
you very likely have money now. 
in all probability they must be 
twenty by this time. 

4. Agarup, "seemingly" 

agarup may at 
agarup madi 
agar&p napigsd 
agarup naimbdg 
agarup masakit 

(preceding, no ligature). Exam- 

he seems to be willing:, 
he seems to be unwilling 
he seems to be strong, 
he seems to be good, 
she seems to be ill. 

5. Komd: used for the conditional, the subjunctive and the 
optative (following, no ligature). Cfr. The Verb, Moods and 
Tenses. V, VI and VII. ^^ 

6. Mdn, kadi, kad: entreating (ordenarily following, no liga- 
ture). Examples: 

sikd man 
mangtidka man 
sidk m4in 
nmdyka man 
napudot man 

you now (inviting), 
please, give me some, 
let me do it. 
come here, please, 
isn't it hot? 

^Philip. Jour. Sci. 75 (1941) 200-202. 

Vanoverbergh: Iloko Adverbs and Prepositions 199 

kas man la dika makagun-od Hi 

paganipemsa man a didk wxiba- 

lin ti agddwat 
kasdi man met Ideng ti inard- 

dikansa man mariknd 
ddkes man ti partdangna 
man met Ideng 
mangdlaka man iti danum 
kitdem kad a siaydt toy ilimi 

dikd kad Ideng mapdn 

kaasidnnak kadi 
tulonganyo kadi 
pakawanem kadi 
ipadymo kadi a kalak-dmanmi 

idawdtannakam kad iti amdm 

innak kad indengdn 

uray kadi ket ti hidg ababd 

di kadi matimano ti tdo a maka^ 

di kad met la sikd ti pangiku- 

anid kad daytoy a maipady iti 

adu a tdo 
anid kad ti kondk idi 
dakdmto kad ket nga addadn 

ndmakto kad a liwliwdda 
indakto kad pay patien 
madayddyaw kadi ti amdm 
sadn kadi apo 
umikkdtka kadi 
anidnto kad ketdi kenkdn a 

bassit ti pammdtina 

Note 32. Man is extensively used in 
between two parts of a complex conjunction, instead of following the 
said conjunction at it stands. Cfr. The Conjunction, passim.40 

you seem not to obtain what you 

do you think perhaps I can't ask 

for things? 
he behaved in exactly the same 

you don't seem to feel it. 
he probably foresees bad news, 
wait a moment (Fll get you), 
please, get water, 
please, look with favor upon our 

don't you go? (you certainly 

please, have mercy on me. 
you should help him, 
please, forgive it. 
please, allow us to partake of 

please, ask your father for us. 
let me go and pay attention to it, 
never mind, life is very short. 

isn't it a shame how few people 

have sense? 
isn't it you to whom I entrust 

what is this for a great number 

of people? 
what did I tell you some time ago? 
how could we have food? (richer 

people haven't), 
great will be their consolation, 
will they believe me? (I doubt it), 
your father should be honored, 
no, sir, please, 
go away, please, 
what is the matter with you 

having so little faith. 

Iloko and is very often placed 

IV. Adverbs implying wonder: 
1. Gdyam (following, no ligature) 

ni dma gdyam it's 

Examples : 

father (I didn't expect 

kastdka gdyam 

that is the kind of man you are 
(I shouldn't have thought it). 

» Unpublished. 


The Philippine Journal of Science 


addd ayd gdyam 

simmangpet gdyam ti andkmo 

isu gdyam 

awdn gdyam 

ni gdyam Ana ditoy 

adddka gdyam ditoy 

is there any, really? 

your son came home (I didn't 
know that). 

it's he (I thought it was some- 
body else). 

there isn't any (I thought there 

well, this is Ann (isn't it 

you are here now (I didn't know 

Giem (following, no ligature; often pronounced diem) 
awdn pay giSm 
dud pay ti gindtangko giem 
awdn met gi4m ti aramidek 
kondna idi gimong apo giem. 

isu ti saona giem 

adu giim ti kUkuana ket di pay 

sadn met giem 
ti komd drak giem apo 

maysd pay ti gatdngek komd 

sagwawalo komd gi4m 
innak komd kumita giim no 

no addd met kukuak giem 

suktdm man iti naingir^pis 

there isn't any yet (I thought the 

I bought two (and still you don't 

I don't do anything to you (why 

are you angry?) 
he said it in the assembly, sir (yes, 

that is what he said, 
he has so much and nevertheless 

he does not give, 
no, not at all. 
better the wine, sir (we prefer the 

I should like to buy one more. 

it should be eight each. 

well, let me go and see if they are 

well, I also have some (and still 

you leave me out), 
change it, please, into a thinner 


V. Adverbs implying resignation: 

1. Anid ngay, anid pay, anid rigarud, "what can be done? 
Examples : 

anid ngay no ndkem ti nagandk 

anid pay no sigud a ddkes 

anid kad pay no napangldw met 

what can be done if it is the will 
of the parents. 

what can be done, he has always 
been bad. 

what can be done, really, as one 
is poor. 
anid ngan^d awdn met sahdli a what can be done, there is no other 



Vanoverbergh: Iloko Adverbs and Prepositions 201 

2. An-anoen {kadi), "what can be done?" Examples: 

an-ano4n no awdn met ti pirdk what can be done if there is no 

an-anoen kadi no isu ti kondna what can be done if that is what 

he says. 


I. Most prepositions are rendered into Iloko by adverbial ex- 
pressions followed by the genitive, and by articles or pronouns 
in the oblique. Examples: 

at midnight, 

before the judge, 
under the tree, 
on the mortar, 
in the jar. 
outside the town, 
outside the house, 
in the middle of the road, 
along the river, 
at the other side of the river, 
east of the post. 
ti between the house and the gra- 
on a line with the old women, 
above the stairs, 
beneath the hill; at the foot of 

the hill, 
behind the man. 
in front of the old man. 
feed it to the dog. 
put it on his head, 
put it in the hole, 
sell it to the girls, 
buy it from Ann. 
tell it to your father, 
give it to him. 
bring it to me. 
take it from us. 
I take it away from this child. 
ibitinmo ltd kayo hang it on that tree. 

II. 1. 'Trom'', indicating the place, is expressed by: 

a. Aggapo (past: naggapo), literally: it comes from; followed 

either by the oblique or by a preposition or an adverb of 

place. Examples : 

aggapo ididy Bontok from Bontoc. 

aggapo sadi Idngit from heaven. 

naggapo ditoy from here. 

iti tengngd ti rabii 

iti ngdlay ti ddlan 

iti sakldng ti okom 

iti sirok ti kdyo 

iti rahdw ti alsong 

iti uneg ti burndy 

iti away ti Hi 

iti rudr ti baldy 

iti tengngd ti ddlan 

iti igid ti karaydn 

iti bdngir ti karaydn 

iti ddya ti adigi 

iti nagbaetdn ti baldy ken 

iti bdtug dagiti baket 
iti ngdto ti agddn 
iti babd ti turod 

iti likuddn ti tdo 
iti masangodnan ti lakdy 
ipakdnmo iti dso 
ikdbilmo iti olona 
iserrekmo iti abUt 
ildkom kadagiti baldsang 
gatangem ken Ana 
ibagdm ken amdm 
itedmo kenkudna 
yegmo kanidk 
aldem kadakami 
ikkatek itoy ubing 


The Philippine Journal of Science 


naggapo ididy 
aggapo ken dvm 
naggapo iti atep 
aggapo ltd baldy 
naggapo itdy lakdy 

from there, 
from my father, 
from the roof, 
from that house, 
from that old man. 

b. The prefixes i or taga joined to the name of the place. 
Cfr. The Adjective. II. Special Forms. S.'*^ Examples: 


he is from Solsona. 
he is from Bintar. 

2. "From", indicating the place or the time, is expressed by : 
a. Mangrugi (past: nangrugi) literally: it begins at; followed 

either by the oblique or by a conjunction, a preposition or an 

adverb. Examples : 

mati^rugi ken Ama 
mangrugi itoy kdyo 
mangrugi ididy karaydn 
mangrugi ididy 
mangrugi iti higdt 
nmngrugi idi kalmdn 
mangrugi no rabii 
mangrugi itd 

from Ann. 
from this tree, 
from the river, 
from there, 
from morning, 
since yesterday, 
from this evening, 
from now. 

b. Manipud (past: nanipud), literally: it begins at; sipud, 
literally: beginning. Both are either followed or not by the 
same terms as the preceding. Examples: 

mnnipud iti karaydn 
manipud kadagiti il-ili 
manipud ken Antonio 
nanipud ididy haldymi 
manipud ditd 
manipud Lawdg 
sipud ditoy 
sipud ken Judna 
mxinipud iti parbangon 
manipud no bigdt 
nanipud itdy bigdt 
manipud itd 
nunipud kinaubingna 
manipud sidm ti bildngen 
sipud idi napdn a tawen 
sipud no bigdt 
sipud itd 
sipud kinaubingna 

from the river, 
from the towns, 
from Anthony, 
from our house, 
from there, 
from Lawag. 
from here, 
from Joan. 

from early in the morning, 
from tomorrow, 
since this morning, 
from now on. 
since his babyhood, 
from the ninth of this month, 
since last year, 
from tomorrow, 
from now. 
since his babyhood. 

«Anthropos U (1931) 483-484. 

Vanoverbergh: Iloko Adverbs and Prepositions 203 

III. "To" or "till", indicating the place or the time, are ex- 
pressed by: 

1. Agpapdn, from the prefix agpa indicating direction and 
the stem pan, "going." Examples: 

agpapdn itd 
agpapdn malem 
agpapdn bigdt 

until now. 
until afternoon, 
till morning. 

2. Agtnngpal or manungpal, literally : it finishes at : followed 
either by the oblique or by a conjunction, a preposition or an 
adverb. Examples : 

to Ann, 
to here, 
to your house, 
till early tomorrow morning. 

agtnngpal ken Ana 
agtungpdl ditoy 
manungpdl ididy baldyyo 
manungpdl no agsdpa 

3. Inggd, literally: end. Inggandy literally: its end, agingd, 
literally: it ends; aging dna (the same as aging gd with an in- 
correct addition of the pronoun na). These prepositions are 
followed or not by the same terms as the preceding. Examples : 

inggd itd 

inggdt malem 

inggdna iti agddn 

inggdna ditoy 

inggdnat iti inggdna 

inggdna ken patdy 

iggdn^t iti tungpdl ti lubong. 

inggdna iti maikapito 

inggdnat no bigdt 

inggdna itd 

inggdnat tungpdl bidgko 

aginggd ken Judn 

aginggd iti maikatlo a bMan 

aginggdt masangodnan 

aginggdnat itd 

till now. 

till afternoon. 

to the ladder. 

to here. 

eternally (literally: till its end). 

till death. 

until the end of the world. 

to the seventh. 

until tomorrow. 

till now. 

till the end of my life. 

to John. 

till the third month. 

till later; until the future. 

till now. 

IV. 1. "With" is ordinarily included in the prefixes maki, 
tagi and ka, in the combination fca . . . an, and in the prefix 
i combined with the prefixes sanga or sangka. Cf r. The sub- 
stantive, III. Formation of Substantives. III. 4; and The 
Verb, The Adjectival Prefix maki; The Adjectival Prefix ag, 
b. Ill B. 1 ; The Substantival Prefix I b. II. Note 6. B. and 
so on.^2 

«Anthropos U (1931) 472-473; Philip. Jour. Sci. 69 (1939) 231, 
235-237; 71 (1940) 20. 

204 The Philippine Jowiml of Science *^^* 

2. Sometimes "with" is expressed by the preposition agra- 
man, followed or not by the oblique, or by the preposition patiy 
followed by the nominative. Examples: 

agrdman kadagiti kakabsdtna with his brothers. 

agrdman kadagiti addlanna with his disciples. 

agrdman kanidk with me. 

agrdman haldy and also the house. 

agrdman anndkna with his children. 

agrdman dmin with all. 

dagiti gayyemko pati ni Ana my friends and also Ann. 

Indiana ti haldy pati dagiti ali- he took the house and also its 

kdmenna furniture. 

ti pirdkmi pati ti pagan-andymi our money and also our clothes. 

V. "Toward" is included in the prefixes agpa and ipa. Cfr. 
The Verb, The Substantival Prefix ipa; Verbs Implying Order 
or Permission. II A. 1.^^ 

VI. Other prepositions are: 

1. Ididy, "at, in, to, into" and so on (in general). Examples: 

addd ididy Lawdg he is at Lawag. 

addd ididy haldy he is in the house. 

simmangpet ididy pagdigHsan he arrived at the bathing place. 

"tnapdn ididy bahd he goes to the lowlands. 

simrik ididy agdmang he entered the granary. 

agydn ididy tdlon it stays in the rice fields. 

immUli ididy atipna he climbed on its roof. 

2. Sadi "in, to," and so on (^exclusively with geopraphical 
names and terms). Examples: 

odtl ti magdtang sadi Manila much can be bought in Manila. 

agnaid sadi Bulakdn he dwells in Bulacan. 

nagddal sadi AmSrika he studied in America. 

inka sadi abagdtan go south. 

napdn sadi amidnan he went north. 

immuli sadi Idngit he ascended to heaven. 

Note 33. Sometimes the name of the place follows another word im- 
mediately without any preposition or article. Examples: 

mapdn Manila he goes to Manila. 

vjapdn Idngit he went to heaven. 

3. Babaen, "through," and so on, followed by the genitive. 
Examples : 

habaSn ti paliibos dagiti agturdy with permission of the authorities. 

hahain ti panangihdhaetna through his mediation. 

babaen ti pannakaihelUnd ti through the shedding of his blood. 


hahain ti panagsublida through their coming back. 

« Philip. Jour. Sci. 71 (1940) 20-22, 26-27. 

Vanoverbergh: Iloko Adverbs and Prepositions 205 

4. Malaksid, "except," followed by the oblique. Examples: 

malaksid iti pusayo except your cat. 

malaksid ken gayySmko except my friend. 

malaksid kadagiti dso except the dogs. 

malaksid kadakayo except you. 

Note 34. Other prepositions, which at the same time may be used as 
conjunctions, will be studied under the Conjunction. 


By Edwin Bingham Copeland 
Of the University of California 


Genus ASPLENIUM Linnaeus 

Brause, Engler's Jahrb. 56 (1920) 144-152, nearly three 
decades ago, listed 49 species in this genus. Two of these are 
Athyrium. One or two others are probably included because of 
wrong identification. He overlooked one published species; five 
have been published in subsequent years; two known elsewhere 
have been found here; and three are here described as new. 
The number to be accounted for would seem to be about 56, of 
which I can distinguish, by comparison or description, about 45. 
In support of the reductions, except as to my own species, the 
isotypes available for comparison are mentioned. 

Fronds simple and entire. 
Nest-forming, with marginal vein. 

Sori broad, long, approaching margin. 

Base gradually narrowed 2. A, Phyllitidis 

Base broad, sessile 3. A, cymhifoliiim 

Sori shorter, narrow, close in full fruit 1. A. Nidus 

Not nest-forming, veins free. 
Sori paired, facing one another. 

Fronds uniform, oblanceolate 46. A. schizocarpiim 

Linear-lanceolate; juvenile fronds com- 
pound .,... 47. A, scolopendropsis 

Sori not paired. 
Frond coriaceous or subcoriaceous. 
Fronds over 25 cm long. 
Apex acute or acuminate. 

Surface squamuliferous 4. A, vittaeforme 

Surface naked. 

Costa prominent 5a, A, aerobryum 

Costa depressed 5. A. comosum 

Apex cuspidate 6. A. Wemeri 

Frond under 25 cm long 

Rigidly coriaceous 7. A, durum 

Subcoriaceous 4a. A, morobense 

* Report on the Third Archbold Expedition, etc. 
8104 6 207 

208 The Philippine Journal of Science ^^* 

Frond herbaceous. 

Rhizome slender, green 8. A» papuanum 

Rhizome stouter, brown 9. A, amboinense 

Fronds pinnate to bipinnatiiid. 

Small ferns with axes dark and polished. 
Pinnae rounded at apex. 

Fronds under 1 cm wide 10. A, Trichomanes 

Fronds broader 11. A, normale 

Pinnae more or less acute. 

Apex of pinnae nearly equal-sided 12. A, unilaterale 

Lower side of pinnae almost wanting- 12a. A. filipes 

Mostly larger, axes not dark and polished. 
Lamina herbaceous, green. 
Pinnae large, not very oblique 

Rachis not winged 13. A. subemarginatum 

Rachis winged upward 13a. A. Regis 

Lower or all pinnae conspicuously oblique. 

Proliferous by slender rhizomes 14. A. bipinnatifidum 

Not proliferous by rhizomes. 
Pinnae under 5 cm long. 

Lower pinnae distant, reduced 15. A. monotis 

Frond truncate at base. 

Pinnae merely toothed 16. A. tenerum 

Pinnae pinnatifid 16a. A. decorum 

Pinnae longer. 

Surface naked 17. A. remotum 

Surface squamulose 18. A. persicifolhim 

Lamina more firm, usually brownish. 
Fronds up to about 5 cm wide. 
Larger pinnae incised. 

Fronds up to 2.5 cm wide 19. A. gracile Fee 

Fronds about 5 cm wide 20. A. planicaule 

Pinnae not lobed. 

Frond compact 21. A. pellucidum 

Frond long and lax 22. A. keysserianum 

Well developed fronds larger. 

Rachis proliferous 23. A. cromwellianum 

Not proliferous. 

Pinnae excised on lower side. 

Rachis more or less scaly 24. A. acutiusculum 

Rachis glabrescent 25. A, falcatum 

Base incompletely excised. 

Pinnae distinctly stalked 26. A. macrophyllum 

Pinnae almost sessile 27. A. Sancti-Christofori 

Fronds bipinnate or more compound. 
Sori dorsal (not marginal). 

Segments linear-cuneiform, 1 mm wide 28. A. insititium 

Segments distinctly wider. 
Rachis bulbiferous. 
Pinnules toothed 31. A. hapalophyllum 

Copeland: Aspleniacem of New Guinea 209 

Larger pinnules subpinnate 29. A. Foersten 

Tripinnate - 30. A. paedigens 

Not proliferous. 

Indusia with parallel sides. 
Pinnules not deeply cut. 

Fronds clustered, thin 40. A. kelelense 

Fronds remote, firm 35. A. settsectum 

Pinnules incised or pinnate. 

Segments cuneate, 2-3 mm wide. 

Rhizome scandent - 36. A, Brassn 

Rhizome short. , , ^ 

Rachis fibriUose 32. A. lanceolatum 

Rachis glabrescent. 

Frond lax ■ 33. A. laxifohum 

Frond compact 34. A. tafanum 

Segments broad. 

Fronds firm to coriaceous. 

Pinnules rounded 37. A. euneatum 

Pinnules subacute 37a. A. affine 

Fronds thin. ^ , .^.-x j* _ 

Bright-green 38. A. laserjnmfohum 

Black-green 39. A. Shawn 

Indusia boat-shaped, ends acute 40. A. nutans 

Sori elongate, straight, free side marginal. ^yhvinnatifidum 

Frond pinnate or -bbipinnate. s„,all « A sn^-^^fi^-^^ 

Fronds decompound, large *"• " 

Sori short, marginal to apical (Loxoscaphe). 

Fronds up to 10 cm long 43. A. Bakeri 

Fronds much larger. novo-guineenBe 

Fertile segments mostly entire *«• ^- "" ,/ , 

Fertile segments mostly furcate 45. A. &c/iM«2ei 

1. A. NKDOS L. 

A. Nidus L., Sp. Plant. (1753) 1079. 
Brass 8966, 11320, 12903, 12951, alt. 20-2,200 m; Carr 1201,0; 
King ^25. 

Old-World Tropics. , 

Questionably distinct from A. Nidus is A. elhpttcum (Fee) 
Copel., Elmer's Leaflets 3 (1910) 819. Referable to it are 
Brass 11373, 12385, 137^2, 13863, 12903; King m; and Clemens 
s n Brass 897U is intermediate between these and what I sup- 
pose to be typical A. Nidus. Brass 137i2 seems to be exactly 
A.nidiforme v.A.v.R.; and Carr 11898 is like it. 
Philippine; Borneo; Java. 


A. Phyllitidis Don, Prod. H. Nepab (1825) 7. 
Brass im7, 13800, Idenburg River, alt. 50 m; King 422, A23, 

210 The Philippine Journal of Science *^^ 

These are unlike typical A. Phyllitidis of India in having 
a veinlet which runs under the free edge of the indusium, and, 
as the latter curls back after maturity, gives superficially the 
same appearance as A, scolopendrioides J. Sm., in the case of 
which this appearance is not due to a vein. There is no such 
vein in typical (?) A. Phyllitidis, and only a suggestion of it 
in A. simonsianum. I suppose that all reports of A. Phyllitidis 
in New Guinea are based on this atypical form. The same 
peculiar vein is found in many collections from the central and 
southern Philippines, including Cuming 819, from Samar. 

3. A. CYMBIFOLIUM Christ. 

A. cymbifolium Christ, Bull. Boiss. II 6 (1906) 999. 

Brass 12076. 


The preceding three species represent PresFs section Tham- 
nopteris. Because all have been construed as forms of A, Nidus, 
and even mixed in collection, their distribution and synonymy 
are uncertain. 


A. vittaeforme Cav., Descr. (1802) 255. 

Brass 11500, Bele River, alt. 2,200 m. 

Both A. vittaeforme and A. squamulatum Blume have been 
reported from New Guinea. As to their probable identity, 
see Christensen, Dansk Bot. Arkiv 9 (1937) 18. To his ob- 
servations, it may be added that I have complete specimens 
from the type region, identified as A. squximulatum, which are 
certainly Cavanilles' species. All Philippine "A. squamulatum" 
is this species, which is also in Borneo and Sumatra. My 
Java specimens so named are distinct, but I suspect wrong 

4a. ASPLENIUM MOROBENSE Copel., sp. nov. Plate 1. 

Micropodium, rhizomate adscendente, 5 mm crasso, paleis 
ovatis acuminatis brunneis usque ad 6 mm longis vestito; 
stipitibus confertis, 10-23 cm longis, gracilibus, paleis parvis 
dense, sed sursum decidue, vestitis; lamina simplice, 15-25 
cm longa, 3.5-4.5 cm lata, acuminata, haud prolifera, basi 
cuneata vix decurrente, integra, subcoriacea, costa paleis paucis 
minutis deciduis praedita, alibi glabra, inferne pallida; venis 
liberis, late patentibus; soris ca % marginem versus elongatis, 
indusiis 0.8 mm latis, sporangiis longissime pedicellatis, annulo 
20-cellulare, sporis oblongo-reniformibus, episporio hyalino, 
haud spinosis. 

Copeland: Aspleniaceas of New Guinea 211 

New Guinea: Morobe, Boana, alt 800-1,300 m. Mary Strong 
Clemens No. il541. 

In the same group and from the same region is A, Werneri 
Ros., which I have not seen. It is described as larger through- 
out, which might well not be a real difference. But it is said 
to be abruptly acuminate, and rounded at the base, while the 
eight fronds of A. morobense in hand are all gradually narrowed 
to both ends. The lamina of A. Werneri is described as scaly; 
the immature frond of A. morobense bears sparse, minute, 
amorphous, ferruginous squamulae on the upper surface, and 
is promptly glabrescent. The nether surface, even as the frond 
grows, is practically glabrous. 

5. A. COMOS13M Christ. 

A. comosum Christ, in Schum. & Lauterb., FL dent. Siidsee (1901) 127. 
I do not know the type, Schlechter 16609 might be authen- 
tic, but its costa is prominent, even carinate on some fronds. 

5a. A. ACROBRYUM Christ. 

A. acrohryum Christ, Nova Guinea 8 (1909) 150. 

Brass 12198, with long, winged stipe; 1S711, with long (10 
cm) terete stipe; 1S81S, attenuate downward and subsessile; 
King 120 A, 89 S and s. n. 

There was considerable confusion in the numbering of King's 
specimens, and, understanding that I had a complete set of his 
later collections (some earlier ones went to Bailey only), I 
suppose that one of these just cited is the plant sent to Buiten- 
zorg and reported as No. 820, the type of A. paucidens v.A.v.R. 
This is a very variable plant, but I believe that all specimens 
cited above are one species. Brass 18188 is identical with 
Schlechter 16609; if that specimen is correctly named, all are 
A. comosum. It is noted that Christ closed his comment on 
A. acrobryum with "Conf. Asplenium comosum Christ de la 
Nouvelle Guinee orientale." 

6. A. WERNERI Ros. 

A, Werneri Ros., Fedde's Repert. 5 (1908) 39. 

Not seen. The type is Werner No. 8, from Mt. Gelu. Re- 
ported also as Ledermann 12516. Reported as with rhizome 5 
cm. thick, short-creeping and then suberect; frond long-stipitate, 
subrounded at base, obtuse and abruptly acuminate. Probably 
near A. vittaeforme. 

212 The Philippine Journal of Science ^^* 

7. A. DURUM Copet Plate 2. 

A. durum CopeL, Univ. Calif. Publ. Bot. 18 (1942) 222. 

Rhizomate breve, valido, paleis ovatis acutis brunneis circa 
3 mm longis vestito; foliis confertissimis, sessilibus, integris, 
lineari-oblanceolatis, usque ad 23 cm longis et 17 mm latis, 
acutis, deorsum longe attenuatis, prope basin inferne parce 
paleatis aliter nudis, rigide coriaceis, costa applanata, venis 
immersis furcatis in marginem cartilagineum plus minus de- 
flexum terminantibus ; soris ad costam et marginem fere attin- 
gentibus, angulo 45° seriatis, indusio lato brunneo crasso. 

Dutch New Guinea: Balim River, alt. 1,600 m, Brass 116S9; 
"occasional on bare limestone on deforested slopes; leaves fleshy." 
The description is of the larger of the two individuals. The 
larger fronds of the smaller one are 7.5 by 1 cm, but fruiting 

Di.stinguishfwl from A. apoense and A. ensiforme by sessile 
fronds; from A. simplicifrons by the position of the sori; more 
rigid and naked than any of these. Such plants as this must 
be supposed tx> be reduced relatives or forms of species of normal 
size for the group, the reduced stature and the texture being 
functions of exposure. Some features of resemblance to another 
local species suggest A. cymbifoliMm, which in full development 
has fronds a meter by 20 cm; but the paleae are different. 

8. A. PAPUANUM Copel. 

A, papuanum CopeL, Philip. Jour, Sci. 6 (1911) 79. 

Known only by the type, King 287. Possibly a juvenile A. 
amboinense, but not to be reduced on the evidence in hand. 

9. A. AMBOINENS£ Willd. 

A. amboinense Willd., Sp. Plant. 5 (1910) 303. 

Brass 11258, 12^86, 12897, 1862$, 18657, 18858, 18668, alt. 700 
to 1,100 m. No. 18657 is aberrant, with subsessile fronds, 
minutely but rather freely squamulose surfaces, and remote 
marginal teeth. 



A. Trichomanes L., Sp. Plant. (1753) 1080. 

A specimen collected by Mrs. Clemens is this species, broadly 

Cosmopolitan New to New Guinea. 

11. A. NORMALE Don 

A. normale Don, Prod. Fl. Nepal. (1825) 7. 

Copeland: Aspleniacex of New Guinea 213 

Brass 12212, 12US7, alt. 1,750 m. For first New Guinea re- 
port, see Christensen, Brittonia 2 (1937) 290, Brass 5451 and 

Asia, Africa, Malaya; reported from Queensland. 


A. iinilaterale Lam., Enc. 2 (1786) 305. 

This comprehensive species is reported from New Guinea, 
but all specimens I have seen are referable to A. filipes, 
Africa; Asia; Polynesia. 

12a. A. FILIPES Cofiel. 

A, filipes CopeL, Philip. Jour. Sci. 3 (1908) 34. 

Brass 18850, alt. 130 m. Collected also by King and by Mrs. 
Clemens. A segregate of A. unilaterale. 


A, subemarginatum Ros., Fedde's Repert, 5 (1908) 372. 
A. Kingii CopeL, Philip. Jour. Sci. Bot. 6 C (1911) 79. 
A. pseudovulcanicum v.A.v.R., Nova Guinea 14 (1924) 8. 

Brass 18859, 18900, alt. 55 and 120 m; Damun, Wernes 68. 

13a. A. REGIS Copel., sp. nov. Plate 3. 

A. rhizomatis apice solummodo viso valido ; stipitibus f ascicu- 
latis, 15-20 cm longis, brunneis, paleis lineari-lanceolatis 2-4 
mm longis, interdum ramis lateralibus ornatis, persistentibus 
vestitis; lamina 30 cm longa, non prolifera, pinnata foliola 
apicale pinnis lateralibus simile, rhachi minute et sparse paleata, 
sursum alata ala 1 mm lata ; pinnis utroque latere circa 6, infimis 
vix diminutis, medialibus 8 cm longis 2 cm latis, acutis, alato- 
pedicellatis basi suboblique late cuneatis, grosse serratis, herba- 
ceis, viridibus, inferne sparse squamulif eris ; venis erecto- 
patentibus, furcatis ; soris linearibus nee costam nee marginem 
attingentibus. Plantae juvenilis stipitibus 6 cm longis, lamina 
jam fertile 12-15 cm longa deorsum bipinnata, rhachi ubique 

Papua : King 880. 

Probably related to A. remotum, as indicated by the bipinnate 
fronds of the juvenile plant. I do not feel altogether sure 
that this is not A. cesatianum Baker, known only by Baker's 
description and by Christensen's reduction to A, persicifolium, 
which this certainly is not. Juvenile plants might have been 
unknown; but Baker could hardly have overlooked the alate 
rachis ; the pinnae are coarsely serrate, not crenulate ; and I find 
no large paleae at the bases of the stipes. 

214 The Philippine Journal of Science ^^^^ 

Isotype in Herb. Univ. Calif.; Lakekamu, King 857 (type 
of A. Kingii), 800. The isotype of A. subs emarginatum and 
the Brass specimens are proliferous at the apex; the apex of 
King's fronds is wanting. The lowest pinnae of King 857, but 
not of No. 800 nor the Brass specimens, are forked; we have 
not the base of Werner 68. The type of A. pseudovulcanicum 
is Lam 671, with Lam 67 A and 7i2 also cited, from the Mam- 
beramo River, about sea-level. I have not seen them, but 
the description shows no distinctive feature. 



A. bipinnatifidum Baker, Synopsis (1867) 221. 

Brass 12881, 12889, alt. 1,200 and 1,500 m. Apparently com- 
mon. The pinnae vary from entire through lobed to subpinnate. 
For synonymy, see Christensen, Dansk Bot. Arkiv 9 (3) (1937) 

To Fiji, and reported in Celebes. 

15. A. MONOTIS Christ 

A. monotis Christ, in Schum. & Lauterb., Nachtr. (1905) 48. 

Known only by the type collection. Described as with small 
sterile and much larger fertile fronds. 

16. A. TENERUM Forster 

A, tenerum Forster, Prod. (1786) 80. 

Brass 8980, Cyclops mountains; 18757, 1U20. Common 
throughout New Guinea. 
Asia to Tahiti. 

16a. A. DECORUM Kaaze 

A. decorum Kunze, Bot. Zeit 6 (1848) 176. 

Darea Belangeri Bory, B^l. Voy. Bot. 2 (1833) 51. 

Asplenium tenerum var. Belangeri of authors. 

Reported from New Guinea as var. acuminata Ros., on the 
basis of Werner 18, also as Ledermann 9298. Pinnae said to 
be up to 12 cm long. Described from the upper part only of 
a frond; very likely not this species. 

17. A. REMOTUM Moote 

A. remotum Moore, Index (1859) 160. 

A. ludens Baker, Joiir. Linn. Soc. 19 (1882) 294. 

Reported from New Guinea are A. dubium Brack., and A. 
multilineatum Hooker, respectively A. remotum Moore and A. 
ludens Baker, all as var. dareoides Ros. 

Copeland: Aspleniacese of New Guinea 215 

First as to the name : Brackenridge described A. dubium and 
A. distansy both applying to the same common Fiji species, 
both names pre-occupied, A. dubium applying to a sterile 
juvenile form with dissected leaves, the fronds of the adult 
plant being simply pinnate. Moore substituted A. remotum 
for A, distans. Then Hooker substituted A. multilineatum 
because he found Moore's name inappropriate. A, ludens was 
described from the Solomons, supposed to be distinguished 
by short sori, perhaps really distinguishable by minute squamules 
instead of narrow paleae of some size on the creeping or even- 
tually scandent rhizome; it, also, has dissected fronds on juvenile 

Following the description of A, multilineatum var. dareoides, 
Rosenstock, Fedde's Repert 10 (1912) 327, proceeds: "Das 
wir es bei ihm mit einer dareoiden Form von A. multilineatum 
Hk. zu tun haben, ergibt sich einmal aus dem charakteristischen 
braunen, kletternden Rhizom mit seinen kleinen, braunen 
Schuppchen, das bei beiden genau iibereinstimmt, sowie zweifellos 
aus einem in meinem Herbar befindllchen Wedel des A. midti- 
lineatum (1. Bamler, Neu-Guinea), der neben den typischen 
ungeteilten Fiedern andere von der Form der var. dareoides 
tragt." We have such a specimen from Rosenstock, 1. Werner, 
bearing three fertile, strictly pinnate fronds, and a number of 
manifestly juvenile dissected fronds. From Alston, we have 
Carr 11677, wholly sterile, but with both dissected and simply 
pinnate fronds, the latter clearly representing a transition from 
juvenile to adult. I have also King 496, 1. Rev. P. C. Shaw from 
mountains behind Taupotu, to which I gave but did not publish 
a new name, distinguishing it from A. remotum by the minutely 
squamulose rhizome, and from A. ludens by sori extending to 
the margin. The latter distinction will probably not hold, and 
the former deserves more observation. These three specimens 
represent one species, ahnost certainly A. ludens, probably also 
A. remotum. Its dissected fronds are juvenile; adult plants 
bear pinnate fertile fronds. As to the variety dareoides, see 
also No. 42, A. scandens. 

Solomon Islands; Fiji, 


A. persicifolium J. Smith: Mett, Asplenium (1859) 97. 
A. cesatianum Baker. Cf. Christensen, Dansk Bot. Arkiv 9 No. 3 
(1937) 46. 

216 The Philippine Journal of Science *^^ 

Collected by Beccari, teste Christensen. 
Philippines; Celebes. 

19. A. €LEMENSIAE Copel., sp. nov. Plate 2 

A. rhizomate breve, paleis fuscis nigrescentibus lanceolatis 
attenuatis vix 1 mm longis vestito; stipitibus approximatis, 
2-3.5 cm longis, brunneis decidue paleolatis; lamina 10-15 cm 
longa, 2-2.5 cm lata, acuminata, basi aut truncata aut paulo 
angustata, subbipinnata, papyracea, bru^nescente, glabra, rhachi 
in parte superiore prolem gerente; pinnis brevi-pedicellatis, 
valde obliquis, majoribus cum pinnula una acroscopica late 
cuneata sessile, parte apicale pinnulae aut incisa aut tantum 
dentata dentibus paucis ; soris secus venas fere omnes elongatis, 
indusio integro persistente. 

New Guinea: Morobe, Boana, alt. 1,000 m. Mary Strong 
Clemens No. A17W. 

This is a reduced member of the group of A, insititium Brack. 

A series of one collection, Copeland 1886, from Mt. Data, 
Luzon, the smallest individual of which matches Fee's description 
and figure perfectly, enables me to identify A. gracile Fee as 
A. planicaule Wall, non Lowe. Similar in appearance as are 
the reduced Luzon and New Guinea specimens, they are not 
immediately related. 

Similar to this, but not identical, is Brass 6^1, — ^see Journal 
Arnold Arboretum 10 (1929) 178. It is more and more finely 
dissected, with less salient venation, about the same size, lamina 
12 by 2.5 cm, and with tiny buds on some rachises. So far as 
the resemblance is due to reduction, it does not prove nor even 
indicate affinity. 

20. A. PLANICAVLE Wall. 

A. planicatde Wall., List. (1828) n. 189 nomen: Mett., asplenium 
(1859) n. 158 non Lowe. 

Reported from New Guinea, but I have no specimen. 
India to Japan and Luzon. 


A. pellueidtim Lam., Enc. 2 (1786) 305. 

Probably common at moderate altitudes. 
African Islands to Polynesia. 


A» Keysserianum Eos., Fedde's Report. 10 (1912) 328. 

Copeland: Aspleniacese of New Guinea 217 

Known only by the type collection, Keysser 26, from Sattel- 
berg; isotype in Herb. Univ. Calif. 


A, cromwellianum Ros. Feddie's Report 10 (1912) 327. 
A, Ledermanni Hieron, Engler's Jahrb. 56 (1920) 150. 

Brass 1124S, 112U, 11598, 12220, 12865, 1SS8S, alt. 850 to 
2,200 m. variable ; pinnae obscurely or sharply toothed, or with a 
basal lobe, but none of this^ collection is as incised throughout as 
is one frond of Keysser 19, the type collection. I have not seen 
A. Ledermanni, but its description does not show that it is 
without the range of these specimens. Moreover, if these were 
Philippine specimens, they would be A. Steerei Harr.; and I 
mistrust identity with A. lobulatum Mett These belong to a 
natural group, within which distinction of species is difficult 
or impossible. 


A, acutiusculum Blume, Enum. (1828) 178* 

Not seen from New Guinea. 

Malaya; Philippines; reported from Samoa. 

25. A. FALCATUM Lam. 

A, falcatum Lam., Enc. 2 (1786) 306. 


Old-World tropics and subtropics. 


A. macrophyllum Swartz, Schroder*s Journal "1800" (1801) 52. 

While distinct enough in typical forms, both this and A. 
falcatum are very variable in New Guinea and Malaya, and the 
assignment of intermediate forms to one or the other is uncer- 
tain. Bakker and Posthumus, Varenflora voor Java 139, treat 
them as one species, and I am now so doing in the Philippines. 
Brass 11638, a dwarf growing on bare limestone, alt. 1,600 m. 
may represent A. macrophyllum. And I would give this name 
to Hollrung 789, distinguished by Hieronymus as A, toyayanum 

African islands to Polynesia. 


A. Sancti'Christofori Christ, in Schum. & Lauterb., Fl. dent. Siidsee 

(1901) 128. 
A. Goadyi Copel. & Shaw, Philip. Journ. Sci. 26 (1926) 330. 

218 ^'/le Philippine Journal of Science *^^ 

Brass 18956, H125, alt. 50 m. Apparently common. 

Endemic in New Guinea and neighboring islands. Goadby 1, 
the type of A. Goadbyi, is from New Britain. 

A. Gjellerupii V.A.V.R., Bull. Jardin Bot. Buit. II No. 7 
(1912) 7, was maintained by its author, Malayan Ferns Suppl. 
I 293, as being A. Sancti-Christofori Ros. non Christ ; but I find 
nothing in the more complete description given there by which 
to distinguish it from the real A. Sancti-Christofori. 

28. A. INSITICUM Brack. 

A. insiticum Barck; U. S. Expl. XVI (1854) 161, pi 22, /. 2 
Brass 1205U, alt. 1,800 m; 12087 (?). Carr 11989 is received 
as A. spathulinum J. Smith, and may be correctly named; but 
I cannot distinguish it from A. insiticium. The two were in- 
cluded in the first description of A, spathulinum by Hooker, 
Sp. Fil. Ill (1860) 170. 

New Caledonia; Fiji; Hawaii; and if A. spathulinum be in- 
cluded, the Philippines. 

29. A. FOEBSTEBI Bos. 

A. Foersteri Ros., Fedde's Repert. 12 (1913) 168. 

Brass 10285, 105U, 10629, 10286, 11242, 1U07, alt. 2,300 to 
2,800 m. The type is Keysesr B 51, isotype in Herb. Univ. 
Calif. Of Brass' collections, the first three are even more 
slender than the type, the lamina up to 50 by 5 cm. The last 
three are much more ample, up to 90 by 27 cm. The first 
three have more promptly glabrous axes ; and are without buds 
on the rachises of the pinnae, as is also our isotype. 


A, cromwellianum, A. insiticium and A. Foersteri are closely 
related species, a group characterized by somewhat fleshy 
texture and the production of gemmae, at least at some of the 
upper axils of the rachis. If A. cromwellianum is A. lohukitum 
and A. insiticium includes A. spathulinum, both species range 
from New Guinea to Luzon (and probably Formosa) and to 
Hawaii. Whether they are one or two species, or a group of 
related species, can only be decided with more ample collections 
than have ever been made. 

30. A. PAEDIGENS Copel. Plate 4. 
A. paedigens Copel., Univ. Calif. Publ. Bot. 18 (1942) 222. 

Euasplenium, rhizomate breve, valido, paleis castaneis acicu- 
lari-lanceolatis integris rigidis circa 3 mm longis vestito; stipite 
25 cm alto 2 nmrt crasso, griseo-fusco, rhachique sparse fibrillosis 

Copeland: Aspleniaceae of New Guinea 219 

mox denudatis ; lamina usque ad 60 cm longa et 25 cm lata, ovata, 
tri-quadri-pinnata, tenuiter papyracea, glabra, apice flagellif orme 
cum pinnis remotis vix 2 mm longis linearibus rhachi gemmif era ; 
pinnis fere sessilibus, apice f rondis simile sed brevius protractis, 
infimis paulo brevioribus, inframedialibus usque ad 17 cm longis 
et 6 cm latis ; pinnulis usque ad 4 cm latis, basi oblique truncatis 
14 mm latis, apice protractis; pinnulisii maximis 8 mm longis 
4 mm latis, ad costam anguste alatam sectis, segmentis 1-3 
cuneiformibus 1 mm latis subacutis vel apice fissis; venulis 
plerumque 2, soris 1 vel 2, linearibus, indusiis laete castaneis 

Dutch New Guinea: Brass 12896, type, 6 kilometers southwest 
of Bernhard Camp, Idenburg River, alt. 1,100 m. "Low epiphyte 
in a rain-forest gulley;'' also, Nos. 12817, ISSSO, 1S752, alt. 850 
to 1,200 m. 

Because of its fine dissection, this has the aspect of a 
Loxoscaphe or Darea, but the sori are attached on the side 
nearest the margin. In spite of the gemmate rachis, this is 
not very close to A. Foersteri, being different in texture as well 
as in dissection. It is not in the group of A. lanceolatum {A. 
furcatuMf A. praemorsum) y in spite of some similarity in 


A. hapalophyllum Ros., Fedde's Repert. 12 (1913) 167. 

Brass 9220, 9295 Lake Habbema, alt. 3,225 m ; Brass & Myer- 
Dress, 10029, Mt. Wilhelmina, alt. 3,560 m; Morobe, alt. 8,000- 
9,000 feet, Clemens s,n. 12S8If bis and the type is Keysser II 
88, and our specimen is marked "unicum." Brass' specimens 
are larger, with a few more residual paleae on the fronds, but 
I am confident of the specific identity. No. 9296, the largest, 
has stipes 20-25 cm long, lamina up to 36 cm long and 15 cm 
wide, tripinnate in the lower part, pinnules with few (2-4) 
secondary pinnules, the lowest sometimes forked. Note on 
the Clemens s.n, specimen reads "pendulous, 4-5 feet." 


32. A. LANCEOLATUM Forsk. 

A. lanceolatum Forsk.: C. Chr., Suppl. Ill (1934) 36. 
A. praemorsum Swartz, Prod. (1788) 130. 

Bamler 29 (1913), from Rook Island, received as "Aspl. 
furcati Thbg. var. an sp. n.", is approximately this "species." 

220 The Philippine Journal of Science ***® 


A. laxi folium V.A.V.R., Nova Guinea 14 (1924) 8. 

Dutch New Guinea: Doorman-top, alt. 2,500 m, Lam 1976. 
Not seen or not recognized. 

34. A. TAFANI7M C. Chr. 

A, tafanum C. Chr., Brittonia 2 (1937) 291. 

Mt. Tafa, alt. 2,400 m, Brass 4839, iS^O. Not seen. 

This and A. laxifolium as described are unlike A, paedigens 
in having stalked pinnae. 

35. A. SE7ISEC11IM Blume 

A. setisectum Blume, Enum. (1828) 187. 

A. inciso'dentatum Eos., Fedde*s Repert. 12 (1913) 167. 

Brass 9297, Lake Habbema, alt. 3,225 m; Clemens 1015i, 
Mt. Sarawaket, alt. 11,000 feet; Keysser B 12, Bolan mountains, 
alt. 3,400-3,800 m, isotype of A. inciso-dentatum in Herb. Univ. 
Calif. The reduction of A. inciso-dentatum is by Backer and 
Posthumus, Varenflora voor Java 339 ; and the fact is not over- 
looked that Christensen, Brittonia 2 (1937) 290, 291, has listed 
them as distinct. 

Malaya; Luzon. 

36. A. BRASSII C. Chr. 

A. Brassii C. Chr., Brittonia 2 (1937) 291. 

Brass 9854, Wilhelmina-top, alt. 3,560 m, epiphytic in large 
masses in subalpine forest. The rhizome, like that of A, seti- 
sectum, is wide-creeping, with remote fronds. 


37. A. CVNEATIJM Lam. 

A. cuneatum Lam., Enc. 2 (1786) 309. 

A. affine Swartz, Schroder's Journal "1800" (1801) 56. 

37a. A. AFFINE Swartz 

These names have been applied rather indiscriminately to spe- 
cimens from Malaya, New Guinea and Polynesia. The original 
distinction between A. cuneatum, from Jamaica, and A, affine, 
from Mauritius, is that the former has rounded, the latter, 
subacute pinnules. 

Brass 18797, 13957, alt. 50 m. Ample, with board, rounded 
pinntiles, A. cuneatum by definition. King 289, 865, smaller 
fronds of the same species. Werner 69, Rosenstock FiL Novog. 

Vtypeland: Aspleniacess of New Guinea 221 

Exsicc. n. 1 is smaller and more harsh, — A. cuneatum var. 
oceanicum Kuhn, King 865 and Bamler, Rosenstock FiL Novog. 
Exsicc. 157 can be A. affine by definition; both rounded and 
subacute pinnules are on both. I am of course not proposing 
the reduction of A, affine. 
Pantropic, as a group. 


A, laserpitiifolimn Lam., Enc. 2 (1786) 310. 

Brass 18969, 140^6, alt. 50 m. Common near sea-level. 
Malaya; Polynesia. 

39. A. SHAWn Copel. 

A. Shawii Copel., Philip. Jour. Sci. 30 (1926) 330, 
Known only by the original collection, 1 SJiaw, King 1^94. 

40. A. KELELENSE Brause 

A. kelelense Brause, Engler's Jahrb. 49 (1912) 29, /. 2 A. 

The original collection, Schlechter 16802, isotype in Herb. 
Univ. Calif., is correctly described as having a short, erect 
rhizome, the fronds fascicled. It looks to me like a juvenile 
plant, possibly of A. cuneatum. Rosenstock Fil. Novog. Exsicc. 
n. 188 y distributed with this name, with a slender, scandent 
rhizome, is no near relative. Six collections by Ledermann, 
det. Brause, Engler's Jahrb. 56 (1920) 146, are unknown to me. 


41. A. NUTANS Ros. 

A. nutans Ros., Fedde's Repert. 12 (1913) 168. 

The type collection, Keysser B 51 A, Bolan mountain, alt. 
2,400-3,000 m, isotype in Herb. Univ. Calif. The indusia look 
like those of a Loxoscaphe, to which it has no other resem- 
blance. Carr 14072, Alola, alt. 6,000 feet, is a more ample 
specimen, with longer and more acute pinnules, more scaly rachis, 
and longer sori, but probably of the same species. 


42. A. SCANDENS J. Smith ^ 

A. scandens J. Smith: Mett., Asplenium (1859) 108. 
A. multilineatum var. dareoides Ros., Fedde's Repert. 10 (1912) 326. 
A, remotum var. dareoides, Engler's Jahrb, 56 (1920) 146. 
A. ludem var. dareoides Alston, Jour. Bot. (1939) 289. 
A. impressivenium v.A.v.R., Bull. Jardin Bot. Buit. II No. XX 
(1915) 8. 

222 The Philippine Journal of Science ^^^ 

As to the variety dareoides, whatever the specific name, 
I have already pointed out in discussing A, remotum, No. 17 
of this enumeration, that the dissected fronds of that species 
represent juvenile stages and are sterile. The variety dareoides 
is a plant with dissected fertile fronds of mature plants. Its 
type was collected by Bamler in the Cromwell mountains in 
1910. We may not have it, but do have a specimen collected 
by Bamler in the same place, dated 1912, Rosenstock Fil Novog. 
Exsicc. n. 152, "Asplenium multilineatum Hk. v. dareoides Ros., 
n. V." It bears two fronds : one sterile, A. scandens, with short, 
rounded pinnules ; the other larger, fertile, with longer subacute 
pinnules. Similar specimens are Schlechter 16428, received as 
A. dubium; Lam 1884, received as A. scandens, and Carr H790, 
15899, received as A, ludens var. dareoides. 

A, scandens of the southern Philippines, — I have 11 collections 
from Mindanao and Palawan, — ^has rather narrow, lanceolate 
pinnae with short pinnules rounded at the apex. My isotype, 
Cuming 297 from Leyte, has longer and more pointed pinnules. 
In New Guinea, the form with narrow pinnae is apparently 
common, represented by Brass 18667, 18846 and 18847. The 
variety dareoides has distinctly longer, more pointed pinnules. 
Brass 12265, 12879 and 12892 are of this form. The extremes 
look distinct, but the type of the species is intermediate. Juve- 
nile plants, identical from Palawan and New Guinea (I. Mrs. 
Clemens) , with fronds down to 5 cm long, are dissected like the 
adults except as necessarily simplified in mechanical correlation 
with stature. 

Philippines; Solomon Islands. 

43. A. BAKEBI C. Chr. 

Known only by the type collection, from Vanapa Valley, alt. 
2,000-4,000 feet. 


A, novo-guineense Ros., Fedde's Repert. 5 (1908) 40. 
Mt. Gelu, alt. 1,000, Werner 16 isotype in Herb. Univ. Calif.; 
Kani mountains, alt. 1,000 m, Schlechter 17191. 

45. A. SCHULTZEI Bratise 

A. Schultzei Brause, Engler's Jahrb. 49 (1912) 30. 
Brass 12022, 12706, alt. 1,800 and 2,000 m; Sattelberg, alt. 
1,500 m, Keysser 196. The type, not seen, is Schuttze (88) 77. 

Vopeland: Asplemacese of New Guinea 223 

Brause, Engler's Jahrb. 56 (1920) 146, lists A. achUleifolium 
(Lam.) C. Chr. as a New Guinea species. Its presence is 

46. A. SCmZOCARPUM Copel. 

A. schizocarpum Copel., Genera (1947) 165. 

Scolopendrium schizocarpum Copel., Philip. Jour. Set. X SuppL 

(1906) 152. 
Diplora schizocarpa C. Chr., Suppl. Ill (1934) 78. 

Schlechter 10229; Bamler, Rosenstock Fit Novoy. Exsicc. n. 
96; Keysser 26; all from Kaiser-Wilhelmsland, alt. ,900 to 
1,600 m. 

Mindanao; very nearly the same form from the Solomon Ids. 


Asplenium scolopendropsis F. v. M., Papuan Plants III (1876) 48, 
Scolopendrium d'Urvillaei Bory, Dup. Voy. Bot. I (1828) 273. 
Diplora d'Urvillaei C. Chr., SuppL III (1934) 78. 
A. Linza Cesati, Kend. Ac. Napoli 16 (1877) 29. 
Triphlebia dimorphophylla Baker, Malesia 3 (1886) 42. 
Scolopendrium Mambare Bailey, Queensland Agric. Journal (1898) 9. 
Phyllitis intermedia V.A.V.R., Bull. Dept. Agric. Ind. Neerl. No. XXI 
(1908) 6. 

It is unnecessary to authenticate this synonymy by citation 
of the many specimens. I am not now in doubt about A. scolo- 
pendropsis; but the Solomon Islands plant to which I gave that 
name, Philip. Jour. Sci. 60 (1936) 114, is nearer to A. schizo- 

Melanesia; Micronesia; Borneo (?). 


Genus BLECHNUM Linnaeus 

Fronds alike, sori not covering nether surface. 

Margin entire 1. B, orientate 

Margin serrate 2. B, nigropaleaeeum 

Fronds dimorphic: Lomaria. 
Fronds pinnatifid or pinnate. 
All sterile pinnae confluent or adnate. 
Dwarfed basal pinnae remote. 

Sterile pinnae over 10 cm long 3. B» saxatile and 

4. B» Ledermanni 

Sterile pinnae up to 8 cm long 5. B. deorso-lobatum 

Dwarfed basal pinnae contiguous, wide.... 6. B. Keysseri 

Only lowest pinnae free, not adnate 7. B. dentatum 

Most pinnae free. 

Dwarfed basal pinnae present 8. B. papuanum i 

Dwarfed basal pinnae absent. 
Pinnae subacute to acuminate. 

8104 7 

224 The Philippine Journal of Science ^^* 

Pinnae entire unless near apex. 
Pinnae rounded at base. 

Pinnae 12 cm long '. 9. B, pendulum 

Pinnae 6 cm long 10. B, Brassii 

Pinnae cordate. 

Apical leaflet like otli^rs 11. B. decorum 

Apical pinnae confluent 12. B, Archboldii 

Margin not entire. 

Pinnae cordate - 11. B, decorum var. 

Pinnae rounded at base. 

C^tae densdy scaly 13. B. Hieronymi 

Costae sparii^ly squamulose - M. B, acutiusculum 

Pinnae obtuse, linear 15. B, revolutum 

Pinnae rounded, oblong 16. B, nudius 

Fronds bipinnatifid bipinnate 17. R Fraseri 


B. orientale L., Sp. Plant. (1753) 1077 (as B. ocddentale). 

Brass 8959, lSi85, alt. 850 m. No. lSi86, with pinna 37 cm 
long and 2.5 cm wide, might be B. finlaysomanum Wall. Com- 

To India, and across Polynesia. 


B. nigropaleaceum Alston, Jour. Bot. 78 (1840) 225; Nova Guinea 
N. S. 4 (1940) 109, PL 4, /. 1. 

Carr HS68, 13S36, Boridi, Papua, alt. 5,000 feet; otherwise 

3. B. SAXATILE Brause 

B. saxatile Brause, Engler's Jahrb. m (1920) 152. 
Ledermann 110 US, Hunstein Peak, alt. 1,350 m. Not seen. 

4. B. LEDEBMANNI Brause 

B. Ledermanni Brause, Engler's Jahrb. 56 (1920) 153. 

Ledermann 11595, 12062, Schraderberg, alt 2,070 m; not 

The seven species (out of eleven credited by him to New 
Guinea) described as new by Brause are presumably distinct, 
and seem to be well described until one undertakes to distinguish 
them by description. 


B, deorso'lobatum Brause, Engler*s Jahrb. 56 (1920) 154. 

Described from Ledermann 11905, 119S2a, Schraderberg, alt. 
2,070 m. To this I refer Brass 10815, alt. 2,750 m; and, less 
confidently, No. 12853, alt. 1,700 m. These are one species, 
but No. 12353 might be B. saxatile. 


Copeland: Aspeneacea of New Guinea 225 

6. B. KEYSSERI Ros. 

B. Keysseri Ros., Fedde's Repert. 12 (1913) 153. 

Keysser 220, Sattelberg Hinterland, alt. 1,400-1,500 m; isotype 
in Herb. Univ. Calif. Well distinguished by the dwarfed basal 
segments, 2-3 mm long by 10-17 mm wide. 

Endemic. , 

7. B. DENTATUM (Kahn) Diels 

B, dpitatum (Kuhn) Diels, in Schum. & Lauterb., Fl. dent. Siidsee 

(1901) 130. 
B. bamlerianum Ros., Fedde's Report. 10 (1912) 325. 

Bamber L. 39, Logaueng, type of B, bamlerianum, not seen. 
Bamber, Rosenstock Fil. Novog. Exsicc. n. 115, Wareo, alt. 600 
m; King 104, Goodenough Bay, alt. 300-600 m. Eelated to 
the Philippine B. egregium, intermediate between Eublechnum 
and Lomaria. 

New Hanover, the type. 

8. B. FAPUANUM Brause 

B. papuanum Brause, Engler's Jahrb. 56 (1920) 158. , 

Lederm^nn 11978, Schraderberg, alt. 2,070 m. Not seen. 

». B. PENDULUM Brause 

B. pendulum Brause, Engler's Jahrb. 56 (1920) 157. 

Ledermunn 11922 Schraderberg, alt. 2,070 m. Not seen. 

1§. B. BRASSII Copel. Plate S. 

B. Brassii Ck)pel., Univ. Calif. Publ. Bot. 18 (1942) 222. 

Lomaria, rhizomate scandente, 4 mm crasso, basique stipitis 
paleis brunneis nonnulis nigrescentibus circa 7 mm longis basi 2 
mm latis incurvis acuminatis vestitis; stipitibus remotis (8 cm), 
primo stramineis, vetustate nigrescentibus, sursum paleis stra- 
mineis plus minus deciduis vestitis, f rondis sterilis 13 cm, f ertilis 
circa 30 cm altiis; lamina circa 30 cm longa, 12 cm lata, basi 
truncata, f oliola apicale maxima usque ad 10 cm longa ; f rondis 
sterilis pinnis inferioribus vix reductis, medialibus 6 cm longis 
11 mm latis, acutis apice obscure serrulatis, subsessilibus basi 
rotundatis, coriaceis, costa infeme decidue squamulata excepta 
glabris; venis gracilibus simplicibus et furcatis obscuris; pinnis 
fertilibus circa 8 cm longis, 2-3 mm latis, superioribus brevio- 
ribus adnatis vix decurrentibus, inlimis brevissime pedicellatis, 

Dutch New Guinea : Idenburg River, alt., 1,800 m, Brass 12287. 
One plant, climbing on dead trunk, in mossy forest. 

226 The Philippine Journal of Science ^^* 

Characterized by the long-scandent rhizome and remote 
fronds, pale-scaly rachis, and complete absence of dwarfed 
basal pinnae. 

II. S. DICORUM Bratise 

B. decorum Brause, Engler's Jahrb. 5^ (1920) 156. 

Ledermann 12i59a, Sepik region, alt. 1,400-1,500 m; also var. 
dilatata, Ledermann 11168; neither seen. To this I refer 
Brass 12iS9, alt 1,800 m; roughly, the distal half of each 
pinna is serrulate. 


U. B. ABCIIBOL0II C. Cliir. 

B. Archboldii C. Chr., Brittonia 2 (1937) 288. 

Brass 4S18, Mt. Albert Edward, alt. 3,680 m, isotyi^ in Herb. 
N. Y. Bot. Garden; Brass 4590, Murray Pass, alt. 2,8^ m. 
Pinnae obtuse to acute; costae clothed beneath with large ovate 


13. B. HIlBON¥MI Brause 

B. Hieronymi Brsmse, Engler's Jahrb. 56^ (1920) 155. 

Ledermann 12217b, Schraderberg, alt. 2,(^0^ m. Not se#n. 
Carr 15182, smaller than described. 

14. B. LATIUSCULUM (v.A.v.B.) C. Chr. 

B. latiusculum (v.A.v.R.) C. Chr., Suppl. Ill (1934) 45. 
Lomaria kitiuscula v.A.v.R., Nova Guinea 14 (1924) 31. 

Lam 1543 Doorman Top, alt. 1,420 m. Not seen. 

15. B. REVOLUTUM (v.A.vJt) C. Chr. 

B, revolutum (v.A.v.R.) C. Chr., Suppl. Ill (1934) 46. 
Lomaria revoluta v.A.v.R., Nova Guinea 14 (1924) 31. 

Lam 1708, Doorman Top, alt 3,250 m; not seen. Brass 9M2 
and 9600, Lake Habbema, alt. 3,225 m, seem to represent this 


16. B. NUDIUS C^pel. I^te 6. 

B. nudim Copel, Univ. Calif. PuW. Bot. 18 (1942) 212. 

B, fluviatile affine eocumque confusum, rhachi sparsissime 
paleacea vel nuda et textura firmiore distin^endum. 

Dutch New Guinea : 7 kilometers north-east of Wilhelmina-top, 
alt. 3,560 m, Brass & Myer-Drees 9966, type, "Sides of moist 
sinkholes and chasms in limestone, common;" also, Brass 9828, 

Copelami: Asplemacem of New Guinea 227 

Lake Habbema, alt* 3,225 m, "Under wet tonks of grassland 
streams; sterile fronds flat-spreading, fertile fronds erect;** 
Brms J^73S, Murray Pass, alt. 2,840 m; Mrs. Clemens s,n, near 
and on Mt. Sarawaket, 9,000 to 13,200 feet. 

Borneo: Mt. Kinabalu, alt. 11,000 feet, Clemens 5H50. Other 
Kinabalu specimens, which I have not, but which are likely to 
be this species, are Clemens 270H, 270 W and 38727. These and 
Brass A733 are determined by Christensen, Gardens' Bull. 7 
(1934) 283, and Brittonia 2 (1937) 288 as J?, ftuviatile (R. 
Br.) Lowe. He states that his Kinabalu specimens (those I do 
not have) conform with those of New 2Jealand in scaliness of 
stipe and rachis. All which I cite from personal observation 
have the rachis conspicuously more naked or wholly so. The 
affinity, however, is to B. fluviatile rather than to the related 
New Zealand species, JB. membranaceum (Col.) Mett., with 
naked rachis. 

The specimens vary greatly in stature, as conditioned by the 
local environment. The type has stipes 15 cm long, laminae 
even 30 cm, the fertile and sterile fronds alike nearly erect. 
Brass 4733 and Mrs. Clemens' collections from Mt- Sarawaket 
are rosette-forming dwarfs growing in open grass. Brass 9328 
and Clemens 51^50 are intermediate. 

17. B. FRASERI (A. Cunn.) Laerssen 

B. Fraseri (A. Cunn.) Luerssen, Flora 59 (1876) 292. 

Brass 108 H, near Lake Habbema, alt. 2,700 m. 

This is var. novoguineensis Brause, Engler's Jahrb. 56 (1920) 
159, distinguished from the New Guinea species by larger si5:e, 
leaves narrowed downward, and strongly toothed segments of 
the sterile pinnae. Only the last feature distinguishes it at all 
from Christ's var. philippinensis, and large specimens of the 
latter are hardly less coarsely toothed, small specimens having 
the subentire segments of the New Zealand type. 

Genus WOODWARDIA J. E. Smith 
W. RADICANS (L.) Smith 

W. radicans (L.) Smith, Mem. Ac Turin 5 (1793) 412. 

Brass 11053, Bele River, alt. probably 2,200 m. "Forest 
undergrowth; common on steep limestone slopes; fronds few, 
in weak spreading clumps.'' 

The genus se^ns to be new to New Guinea. The extension 
of range is significant, because Woodvjardia, always regarded 
as a relative of Blechnum, is a genus of distinctly northern 

228 TAe Philippine Journal of Science ^^* 

distribution, previously known south of the equator in Java 
only, while its relatives are of evident antarctic origin. The 
new collection carries the genus back along the track along 
which the great mass of oriental ferns seems to have immigrated. 
As to the specific name, the recent tendency, — see Christensen, 
Suppl. Ill 196, — ^is to distinguish specific segregates long re- 
garded as forms of W* radicans, among these, W. orientalis, 
W. aurictdata and W. unigemmata. It is possible to distinguish 
W. orientalis and W. unigemmata; but I am not so sure that 
the latter is not W. auriculata, and so prefer for the present 
to use the most inclusive name. Woodwardia is common in the 
mountains of northern Luzon, and unknown between those 
mountains and New Guinea. With the material in hand, it is 
impossible to distinguish the plants of the two regions. 


PertOe pinnae rolled back 1. S. areolaris 

Fertile pinnae Hat. 

Fertile pinnae 2-3 nam wide 2. S, palustris 

Fertile pinnae at least 5 mm wide 3. 5. juglandifolia 

1. S. AREOLARIS (Harr.) Copcl. 

S. areolaris (Harr.) CopeL, Philip, Jour. Sci. 2 C (1908) 406. 

Bra^s 1BU17, alt. 850 m, on Pandanus; Clemens s. n, Morobe, 
alt. 1,300 m, on banana and betel. The latter is the first 
record of the species as growing elsewhere than on Pandanus. 
Other New Guinea records by Holttum are Versteeg 1102 and 
Docters van Leeuwen 9909. 


2. S. PALUSTRIS CBmrm.) Bt4d. 

S. palustris (Burm.) Bedd., Ferns Brit, India Snppl. (1876) 26. 

Brass 1S9S8, alt. 50 m; Brass 9S6, Carr 11323, Papua, sea 
To India and Samoa. 


S. juglandifolia Fxesl, Epim. (1849) 164. 

Bamber 112p, Sattelberg; King 167, Papua; Bamber R, 1, 
from Ins. Rook. 

Philippines; Solomon Islands. 

As to the distinctness of this species, see Holttum, Gardens* 
Bull. 9 (1987) 139. Recognized as a species, 5. juglandifolia is 
the typ^species of the genus. 


[All are Photographs of the Types) 

Plate 1. Asplenium morobense Copel. 

2. A, durum Copel. 

A. Cle'fnensiae Copel. 

3. A. Regis Copel. 

4. A. paedigens Copel. 

5. Blechnum Brassii Copel. 

6. B, nudius Copel. 


Copeland: AsPLEHiAciJS OF New Guinea. | 

fPii 11.11*. JtHTft, BrL, T8, No., 2. 



iiMniU'. JoijB. SCL, 78, No. 2. 


Vmn-ihAKi): Asi'i.K.\'i,\«i-:,K m-' Nkw <U'ixka.| 

[Philip. Jtiru. Sn., 78, No^ 2. 



'#i,\ . . ^ •>^' 


Copelamd: ASPL1NIAC1J3 OF New Guimea.] 

[Philip. Joui, Sci., 78, No. 2. 



J^'^ I 

f«b . '•"•'" 



[Philip. Jour. Sci., 78, No., 2. 





Books reviewed here have been received by the Philippine 
Journal of Science from time to time and acknowledged in this 


Radio Aids to Navigation. By R. A. Smith. Cambri4ge, At the University 
Press, New York, The Macmillan Company, 1948. 114 pp. Price, 

The material for this book was originally prepared as a 
contribution to the British Ministry of Supply (Air) Scientific 
War Records, but by permission of that ministry, it was later 
revised for publication. 

This book traces the remarkable development in the applica- 
tion of electronics to navigation during the war and shows how 
the limitations of prewar radio navigation systems were over- 
come by the introduction of new techniques particularly by the 
development of radar. These electronic devices are now widely 
adapted for air and sea navigation and other works requiring 
quick and accurate determination of position under any weather 
condition. Other devices find application in the field of survey- 
ing, triangulation and aerial photography. 

The book is a good reading matter for the layman interested 
in the sciences, and to those possessing the basic knowledge of 
radio this book ushers them into the relatively new field of 
electronics. — L. G. A. 

Analytic Geometry. Revised edition. By Roscoe Woods. New York, The 
Macmillan Company, 1948. 322 pp. Price, |3.50. 

Beginners on analytic geometry will find this book neither 
too technical nor too elementary because the author introduced 
each new principle with illustrating examples completely 
worked out in the text. Of course, there is the usual dose 
of exercises which are carefully graded at the end of each 
chapter. All the materials needed in the usual short course 
in analytic geometry are presented in the first ten chapters; 
the last five chapters make it fuller and rounder, aside from 
thoroughly introducing the student to solid geometry and the 
concepts needed in the study of calculus. The author stepped 
farther from the usual run of freehand drawings when he 


232 The Philippine Journal of Science **** 

utilized pictures of wire models to represent quadric surfaces 
in chapter XV. Answers to the odd numbered exercises only- 
are given at the back of the book but the author will furnish 
upon request an answer-book with answers to even-numbered 
exercises. — ^L. G. A. 

Strength of Materials. By Joseph Marin. New York, The Macmillan 
Company, 1948. 464 pp. iUus. Price, |4.75. 

The designing of structure and machine members is the object 
of this book. It emphasizes that materials for construction 
must be simple and cheap yet strong enough to resist the dif- 
ferent kinds of stresses be it bending, compression, twist, stretch 
or shear. It calculates to the minutest detail the kind of stress 
that is required in the different parts of the structure and 
machine. The materials that are to compose the structure and 
machines are to be selected and their strength, calculated in 
order to enable th^ai to withstand the shock or stresses from 
within as well as from without. 

This book is a good text and reference material. It is highly 
reconmiended for engineering students, engineering professors 
and practicing engineers. — ^R. R, O. 

The Philippine 

Vol. 78 SEPTEMBER, 1949 No. 3 


Of the one-hundred fourteen charter members originally 
appointed by the Governor-General of the Philippines in 1934 
following the enactment by the Philippine Legislature of Act 
No. 4120 creating the National Research Council of the Philip- 
pines, only eighty-one remained at the time of the death on 
August 6, 1949 of the late Jose Kabigting Santos, former 
Professor and Head of the Department of Botany of the 
University of the Philippines. This irreparable loss of one of 
the outstanding Filipino scientists was keenly felt by his many 
friends and associates. The National Research Council of the 
Philippines, the consulting and advisory body to the Republic 
on matters pertaining to science and scientific research, has 
been directly affected by the death of Doctor Santos, not only 
because of the passing of the charter member who was re- 
sponsible for the idea of holding the first Philippine Science 
Convention in 1932 during his incumbency as President of the 
Philippine Scientific Society, but also because of the marked 
decrease in the ranks of its members devoted to botanical 
research. Thus Dr. Bienvenido M. Gonzalez, President of the 
University of the Philippines, Charter Member and for more 
than ten years, Chairman of the National Research Council of 
the Philippines, fittingly eulogized the late Jose Kabigting Santos 
both in the obituary prepared by the Executive Board of the 
National Research Council of the Philippines and in the an- 
nouncement of the passing of Jose Kabigting Santos transmitted 
to every member and associate of the National Research Council 
on August 11, 1949. 

Jose Kabigting Santos was born in Arayat, Pampanga, on 
February 4, 1889. The son of Lucio de los Santos and Natalia 
Kabigting, he was at the time of his death a little over 60 years 
954a 275 . 

276 The Philippine Journal of Science ^^' 

of age. His early education was obtained in the Liceo de 
Manila, one of the first private schools founded and supported 
by Filipino educators from whom a number of undergraduate 
and advanced students were selected to become members of the 
first group of Filipino pensionados who were sent by the 
Philippine Government to various colleges and universities of 
the United States to pursue advanced professional, scientific 
and technical studies. At the beginning of the academic year 
1911-1912, Jose Kabigting Santos enrolled in the Course in 
Pharmacy offered in the College of Liberal Arts, which was to 
become the College of Pharmacy of the University of the Phil- 
ippines with the late Dean Andrew Grover DuMez of the Col* 
lege of Pharmacy of the University of Maryland as its first 
director. Even during his residence as a student in the Uni- 
versity of the Philippines, he displayed a keen interest in botany 
and an exceptional ability in scientific illustrations especially 
in the different courses he took in the Department of Botany 
of the University. He also showed enthusiasm in other activ- 
ities that the state's highest institution of learning was promot- 
ing among her students. For his ability and leadership, the 
student body of the School elected him President of the Univer- 
sity of the Philippines Pharmaceutical Association. This or- 
ganization included among its members every student enrolled 
in the School of Pharmacy and was undertaking activities 
designed to promote the welfare of pharmacy students and 
the success of the School of Pharmacy in its avowed mission. 

Serving as Student Assistant in the Department of Botany 
of the University of the Philippines, he graduated in 1916 with 
the degree of Graduate in Pharmacy. Immediately after his 
graduation, he was appointed Assistant Instructor in Botany 
in the College of Liberal Arts. In 1918, he was promoted to the 
position of Instructor in Botany. During the early years of his 
connection with the Department of Botany of the College of 
Liberal Arts, he had the opportunity of associating himself 
with well-known botanists who had spent the greater part of 
their lives undertaking botanical research in the Philippines, 
such as Dr. Elmer D. Merrill, former professor of botany in the 
University of the Philippines and Director of the former Bureau 
of Science, now with Harvard University; and the late Dr. 
William H. Brown, former professor of botany in the University 
of the Philippines and Director of the former Bureau of Science. 

Realizing his bright prospects, the University authorities 
appointed him Fellow of the University of the Philippines 

^^' ^ Jose Kabigting Santos 277 

abroad. He left Manila in August, 1919, together with other 
pensionados of the Philippine Government. Upon his arrival 
in the United States, he enrolled in the University of Chicago. 
Notwithstanding his handicaps for not having completed a 
bachelor's degree, he was allowed to enroll in the University of 
Chicago in courses that would prepare him for graduate work. 
The excellent work he performed enabled him to obtain in 1921 
the degree of Bachelor of Science with honors from the 
University of Chicago. He then enrolled in the Graduate 
School of the same university, majoring in botany. At the end 
of the academic year of 1922, the same university conferred upon 
him the degree of Master of Science. He continued his 
graduate work leading to the doctorate degree. At the com- 
mencement exercises of the University of Chicago in 1923, he 
was conferred the coveted degree of Doctor of Philosophy magna 
cum laude. His main research work, the results of which he 
used for his doctoral dissertation, dealt mainly with his studies 
on the differentiation of chromosomes and determination of sex 
in Elodea. These two papers were published in the Botanical 

While in the United States, Jose Kabigting Santos and Dr. 
Joaquin M. Maraiion were appointed official delegates of the 
College of Pharmacy of the University of the Philippines to the 
1921 Annual Meeting of the American Association of Colleges 
of Pharmacy, formerly American Conference of Pharmaceutical 
Faculties held in Cleveland, Ohio. They also represented the 
Philippines in the 1921 Annual Meeting and National Conven- 
tion of the American Pharmaceutical Association held also in 
Cleveland, Ohio. On his way home in 1923, he had occasion 
to visit leading botanical laboratories in renowned universities 
in London, Vienna, Paris, Belgium, and Switzerland. 

Upon his return to the University of the Philippines, he was 
appointed Assistant Professor of Botany in 1924 in the College 
of Liberal Arts. He devoted his time to teaching different 
botanical courses in the University of the Philippines. There 
was not a single year when he did not conduct and supervise 
the botanical excursions that the Department of Botany 
organized regularly for the purpose of instilling in the minds 
of students of botany the importance of systematic botany 
and the indispensable corollary of this work, mainly the actual 
work in the field that enable systematic botanists to learn plant 
characteristics. He also began studies in the pharmacognosy 
of Philippine medicinal plants as early as the first year after 

278 The Philippine Journal of Science ^^^ 

his arrival in the Philippines from his trip abroad. His first 
contribution to the pharmacognosy of Philippine plants was a 
paper on Chenopodium ambrosioides L. published in the Phil- 
ippine Journal of Science in 1925. In 1925 he was appointed 
Acting Head of the Department of Botany of the University of 
the Philippines. In 1926 he was promoted to Associate Prof- 
essor of Botany. 

During all the years- of his teaching of both undergraduate 
and graduate courses in botany, he was actively engaged in 
undertaking research on cytology and the pharmacognosy of 
Philippine medicinal plants. It was especially his appointment 
as part-time Botanist of the former Bureau of Science in 1924 (a 
position which he held continuously for ten years until 1934) 
that enabled him to devote a great deal of his time to botanical 
research. He was also invited to contribute a paper on cyto- 
logy in a special jubilee number of Cytologia issued by the 
International Cytological Association with headquarters in Jap- 
an. Not only was he an active research worker but he also 
spared no efforts in attracting many of his young associates 
to study advanced courses in botany and to undertake original 
botanical investigations. He did this work so well that he was 
instrumental in sending many young members of the faculty 
to pursue advanced studies in the United States. Several of 
those students have returned to the Philippines to serve their 
country and people after obtaining their advanced training in 
various phases of botanical work. 

In 1934, he became permanent Head of the Department of 
Botany of the College of Liberal Arts of the University of the 
Philippines. When an exchange of professors between the 
University of Michigan in Ann Arbor and the University of 
the Philippines was established in 1934, Professor H. H. Bartlett, 
Chairman of the Department of Botany of the University of 
Michigan, came to the University of the Philippines as Exchange 
Professor. At the same time the late Jose Kabigting Santos 
was appointed Exchange Professor of Botany in the University 
of Michigan. There he spent the academic year 1934-1935 
teaching botany. During this second trip abroad, he attended 
the Sixth International Botanical Congress held in Amsterdam, 
Holland, as an official delegate of the University of the Philip- 
pines. On his way home from Europe, he and his family visited 
London, Berlin, Brussels, Belgium, and France. They also spent 
three months in Spain, partly because they were stranded in 

^*» 3 Jose Kabigting Santos 279 

Europe as a result of the Italo-Ethiopian War. From Spain 
they went to Italy and then to Marseilles, France, where they 
finally took passage for Manila. 

As president of the Philippine Scientific Society, then a very 
active instrument for the dissemination of scientific knowledge, 
he was mainly responsible for the idea of holding the First 
Philippine Science Convention in 1932. Under his leadership, 
this national science convention took shape. 

Scientists from different branches of the Government, the 
University of the Philippines, and other universities, colleges 
and professional organizations cooperated wholeheartedly to 
make that memorable convention a huge success. It was dur- 
ing that convention when at the behest of Filipino scientists, 
the late President Manuel L. Quezon publicly announced his 
support for the establishment of a National Research Council 
of the Philippines. Thus Jose Kabigting Santos contributed 
his share in more ways than one to the progress of science in 
our country. 

The full appreciation of the value of research by Jose Ka- 
bigting Santos is well illustrated by the following excerpts 
from his presidential address entitled "Science and the Country" 
delivered during the Opening Session of the First Philippine 
Science Convention : * 

As I have stated before, scientific mindedness is necessary for the 
advancement and prosperity of this country; but scientific mindedness 
alone without its necessary tool — research — ^is meaningless — an empty 
phrase! And I need not tell you that for the development of our natural 
resources, research in pure and applied sciences is indispensable; nor do 
I need to tell you that our country is economically poorer, though potentially 
richer, than many countries of its size in the world because we are behind 
in scientific research. It is high time, therefore, that we give our most 
serious attention to this vital question. We must solve it now; further 
dela^ may prove disastrous to our economic future and fatal to our 
political aspirations. Our government must establish more experimental 
stations, and improve upon and enlarge its existing laboratories, or 
authorize the establishment of new ones. Our colleges and universities 
must within the limit of their ability and resources, do their full share 
in furthering scientific research. In fact, they should be made responsible 
for the scientific progress of the country inasmuch as they are responsible 
for the scientific education of our youth. 

* Proceedings of the First Philippine Science Convention held under 
the auspices of the Philippine Scientific Society, March 15, 16, 17 
inclusive, 1932, Manila. University of the Philippines Natural and Applied 
Science Bulletin, v. 2:165-308. 

280 The Philippine Journal of Science *^* 

We, who have had the fortune of being trained in the ways of science, 
who live in intimate contact with it, who labor under its lure and influence, 
have a duty to perform and a responsibility to discharge towards our 
country and people. We do not, we cannot, dodge this duty and this 
respo^jsibility; but to faithfully fulfill them we need help and encourage- 
ment. We have faith in our ability to build the scientific structure of 
our country provided adequate means are given to us. As the frame- 
work of this structure it would perhaps be necessary to organize an 
institution like the National Research Councils or the National Academy 
of Sciences of various progressive countries. 

With the ever living, ever growing desire in our hearts to serve this 
Pearl of the Orient Seas — ^the one spot in the whole creation which God, 
in His infinite wisdom, has given us to dwell in happily, to own, to 
develop for our benefit and contentment, to love and to defend — ^let us 
exert our best effort to make her self-sufficient and promote her general 
welfare by leading her along upon the charted highway of science and by 
inculcating in her youth the love of research for the sake of truth and for 
the material blessings that research affords. 

Speaking of Doctor Santos, the President of the University of 
the Philippines, Dr. Bienvenido M. Gonzalez, Chairman of the 
National Research Council of the Philippines, said: 

As President of the Philippine Scientific Society, he was mainly 
responsible during his incumbency of introducing the idea of holding a 
Philippine Science Convention in 1932. It was such a success that when 
the National Research Council of the Philippines was created, the 
Philippine Science Convention was held regularly under the joint auspices 
of the National Research Council of the Philippines and the Philippine 
Scientific Society until the war broke out. 

As an author, he published a manual entitled, Experimental 
Botany (Manila, Educational Supply, 1934, pages XIV, 235). 
This manual is at present used by the University of the Phil- 
ippines in the classes of botany in Quezon City, Cebu and Iloilo 
and by other universities and colleges in the Philippines. He 
was contributing editor of The Philippine Journal of Science and 
the University of the Philippines Natural and Applied Science 

He held membership in professional and learned societies, 
namely, the Philippine Pharmaceutical Association, the Phil- 
ippine Scientific Society, the Colegio Medico-Farmaceutico de 
Filipinos, the American Association for the Advancement of 
Science, of which he was a Fellow, the Sigma Xi and the Society 
for Advancement of Research. In the National Research 
Council of the Philippines he held membership in the Sections of 
Pharmacology, Organic and Plant Chemistry, Pharmacopoeia, 
Genetics, Botany, and Chemical and Biological Defense. He 
is listed in the American Men of Science, 4th edition, 1927, New 

^^' 3 jQse Kabigting Santos 281 

York, the Science Press, p* 854, in the World Biogi'aphy, 1948, 
published by the Institute for Research in Biography, 296 
Broadway, New York 7, New York; and also in Bulletin No, 
7 issued in February, 1934, of the National Research Council 
of the Philippines, wherein biographical data and a bibliography 
of his works are given (pages 734-736). 

On September 30, 1925, he married Felisa de la Peiia, a 
holder of the degree of Pharmaceutical Chemist from the 
College of Pharmacy of the University of the Philippines with 
whom he had three children. 

His love of pharmacy manifested itself again when during 
the latter part of his life he and his wife engaged in the 
manufacture in their own laboratory of the well known JofePs * 
preparations, namely Jofel's medicinal cigarette, JofeFs asthma 
relief powder and Jofel's asthma paper. These preparations 
were made chiefly from the flowering tops of Datura alba Nees, 
or Datura fastuosa Linnaeus and from the entire herb of 
Euphorbia hirta Linnaeus. 

The appended bibliography represents the results of the pains- 
taking research work that he pursued during his life. 


1. Differentiation among Chromosomes in Elodea. Botanical Gazette 

75 (1923) 42-59. 

2. Determination of Sex in Elodea. Botanical Gazette 77 (1924) 353-376. 

3. A Pharmacognostical Study of Chenopodium ambrosioides Linnaeus 

from the Philippines. Philip. Jour. Sci. 28 (1925) 529-547. 

4. Histological Study of the Bark of Alstonia scholaris R. Brown from 

the Philippines. Philip. Jour. Sci. 31 (1926) 415-429. 

5. Pharmacognostical Study of Datura alba Nees and Datura fastuosa 

Linnaeus from the Philippines. Philip. Jour. Sci. 32 (1927) 267-296. 

6. Stem and Leaf Structures of Tinospora rumphii Boerlage and Tinos- 

pora reticulata Miers. Philip. Jour. Sci. 35 (1928) 187-208. 

7. A Cytological Study of Cocos nucifera Linnieus. Philip. Jour. Sci. 

37 (1928) 417-437. 

8. Histological and Microchemical Studies on the Bark and Leaf of 

Artabotrys simveolens Blume from the Philippines. Philip. Jour. 
Sci. 38 (1929) 269-282. 

9. Leaf and Bark Structure of Some Cinnamon Trees with Special 

Reference to the Philippine Species. Philip. Jour. Sci. 43 (1930) 

10. A Critical Morphological Study of Thalassia hemprichii (Ehrenb.) 

Aschers from the Philippines. Univ. Philippines Nat. and Appl. Sci. 
Bull. 1 (1930) 1-19. 

♦ The word was coined from the first syllable of Jose and Felisa, his 
first name and that of his wife. 

282 The Philippine Journal of Science *^^ 

11. Pharmacognosy of Philippine Drugs. Jour. Philippine Pharm. Assoc. 2 

(1930) 148-162. 

12. Philippine Drugs. Jour. Philippine Pharm. Assoc. 2 (1930) 211-212c, 

245~249a, 282-287; 3 (1931) 124-129, 189-194; 4 (1932) 316-327, 

13. Anomalous Stem Structure in Archangelisia flava and Anamirta 

cocculus from the Philippines. Philip. Jour. Sci. 44 (1931) 385-407. 

14. Leaf and Seed Structure of a Philippine Coriaria. Philip. Jour. Sci. 

46 (1931) 257-268. 

15. A Contribution on the Life History of Dendrobium anosmum Lindley, 

by Maria D. Pastrana and Jose K. Santos. Univ. Philippines Nat. 
and Appl. Sci. Bull. 1 (1931) 133-144. 

16. Anatomical Study on the Culm of Five Philippine Bamboos, by 

Gregorio T. Velasquez and Jose K. Santos. Univ. Philippines Nat. 
and Appl. Sci. Bull. 1 (1931) 281-318. 

17. Scientific Contributions of the Department of Botany. Univ. Philippines 

Nat. and Appl. Sci. Bull. 1 (1931) 405-413. 

18. The Laticiferous Vessels and Other Anatomical Structures of 

Excoecaria agtallocha, Philip. Jour. Sci. 47 (1932) 295-304. 

19. Morphological and Chemical Studies on the Seeds of Erythrina 

variegata var. 'orientalis (Linnaeus) Merrill, by Joaquin Maranon 
and Jose K. Santos. Philip. Jour. Sci. 48 (1932) 563-580. Also 
in Univ. Philippines Nat. and Appl. Sci. Bull. 2 (1932) 215. 

20. On the Microsporogenesis of Sacchamm spontaneum Linnaeus with 

Special Refenrence to Its Chromosome Number. Univ. Philippines 
Nat. and Appl. Sci. Bull. 2 (1932) 211. 

21. Science and the Country. — Address by Dr. Jose K. Santos, President of 

the Philippine Scientific Society, during the First Philippine Science 
Convention held under the auspices of the Society, March 15-17, 
1932, in the Univesity of the Philippines, Manila. Univ. Philippines 
Nat. and Appl. Sci. Bull. 2 (1932) 174-180. 

22. Morphology of the Flower and Mature Grain of Philippine Rice. 

Philip. Jour. Sci. 52 (1933) 475-497. 

23. Macrosporogenesis of Ldlium philippinenss Baker. Paper read at 

the Fourth Philippine Science Convention held under the auspices 
of the National Research Council of the Philippines and the Phil- 
ippine Scientific Society, February 23-27, 1937, in the University 
of the Philippines, Manila. Abs. Nat. Research Council of the 
Philippines Bull. No. 19 1938) 105. 

24. Anatomical and Blicrochemical Studies of Stropfmnthus cumingii A. 

DC. Paper read at the Fourth Philippine Science Convention held 
under the auspices of the National Research Council of the Philip- 
pines and the Philippine Scientific Society, February 23-27, 1937, in 
the University of the Philippines, Manila. Abs. Nat. Research 
Council of the Philippines Bull. No. 19 (1938) 137-138. 

25. A Tentative Pharmacopoeial Monograph on Tiki-tiki <or Rice Bran. 

Revista Filipina Med. Farm, 28 (1937) 337-342. 

26. The Need for an Extensive Investigation of Philippine Medicinal Plants. 

Revista Filipino Med. Farm. 29 (1938) 165-168. 

27. A Pharmacognostical Study of Mallotus philippinensis (Lam.) MiielL- 

Arg. Revista Filipina Med, Farm, 29 (1938) 405-416. 

^«» 3 jQg4 Kabigting Santos 283 

28. Pharmacognostical Study of Strophantus cumingii A. DC. from the 

Philippines. Revista Filipina Med. Farm. 30 (1939) 365-386. 

29. The Origin and Development of the Female Gametophyte of Hedychium 

coronarium Koenig. Paper read at the Fifth Philippine Science 
Convention held under the auspices of the National Kesearch Council 
of the Philippines and the Philippine Scientific Society, February 
21-26, 1939, in the University of the Philippines, Manila. Abs. 
Nat. Research Council of the Philippines Bull. No. 32 (1939) 

30. Macrosporogenesis and Development of the Female Gametophyte of 

Asparagus sprengerL Paper read at the Fifth Philippine Science 
Convention held under the auspices of the National Research 
Council of the Philippines and the Philippine Scientific Society, 
February 21-26, 1939, in the University of the Philippines, Manila. 
Abs. Nat. Research Council of the Philippines Bull. No. 23 (1939) 

31. Pharmacognostical Study of Quisqualis indica, Lochnera rosea, 

Euphorbia hirta and Premna adorata, Revista Filipina Med, Farm. 
32 (1941) 197-212. 

Patrocinio Valenzuela 
Dean, College of Pharmacy 
University of the Philippines 




By Leopoldo B. Uichanco 
College of Agriculture, University of the Philippines 

Ectenus Dallas is a small genus, which so far has been 
recorded almost exclusively from the Philippines, with the 
exception of a single species from New Guinea (Waigiu and 
Mysol). Hence, although the group is considered a Papuan 
derivative (Breddin, Abhandl. Naturf. Ges., Halle 24 [1901] 
164), its headquarters apparently lie in the Philippines, wherein 
each species exhibits a strong local endemism. The known 
forms may be differentiated with the aid of the key included 
in the present paper, although in some cases separation is 
rather difficult, owing to the tendency of certain species to 
merge. The habits of none of the species are known. Unfor- 
tunately, because there was no material of pvdictis and 
brevirostris on hand, sole reliance had to be made on published 
descriptions for their apparent diagnostic characters. All the 
specimens referred to herein, which formed part of the collec- 
tions of the Bureau of Science and of the College of Agriculture, 
were destroyed by enemy action in February, 1945. It will 
probably take years to reconstitute the collection. However, 
it is deemed that the present diagnostic treatment will help in 
the recognition of the various forms, so that, although a restudy 
is indicated based on new material, release of this report 
obviously has its justification. Moreover, the specific collection 
localities given for each form serve to supply these distributional 
data for papers by Stal and other earlier authors, who merely 
reported "Philippines" or "Manila" for material studied by 
them from any of the islands of the Philippine group. 

The species included in the present paper are: 

1. Ectenus generostts Stal 

Sulu Archipelago 

2. E. mesoleucus Bergroth 

Luzon and Polillo 

1 Experiment Station, College of Agriculture, Contribution No. 1543. 
Received for publication August 5, 1949. 


286 The Philippine Journal of Science *^^ 

3. E. spinosus sp. nov. 


4. E. pudicus Stal 

Mysol and Waigiu 

5. E. spectabilis (Burmeister) 


6. E. SBvem Stal 

Biliran, Bohol, and Mindanao 

7. E. brevirostris Stal 

Philippines; specific locality unknown 

Genus ECTENUS Dallas (1851) 

Genotype: Cimex spectabilis Burmeister (Philippines) 

Close to Dalpada Amyot et Serville (tribe Halyiini), differing 
therefrom principally in that the first antennal segment in 
Ectenus is longer than the marginal space on the head between 
the eye and the insertion of the antenna, the first rostral segment 
extends markedly beyond the buccute, the part of the scutellum 
bounded by the frena is much longer than broad at the base, 
and the postfrenal part is narrower. 

Key to the known species of Ectenus 

1. Bucculae not prominently produced anteroventrad, anteriorly de^ 

scribing a right or subacute angle. Postfrenal part of scutellum 
broader, more or less broadly rounded apically, almost entirely 
stramineous, sublsevigate, sparsely punctured; basal angles each 
with a stramineous, sublsevigate spot, which is as large as apical 
spot or smaller. Robust species; width of body about one-half 

length 2 

Bucculae prominently produced anteroventrad into an acute or subacute 
process. Postfrenal part of scutellum narrower, roundly subacute 
at apex; stramineous, laevigate spot only thinly bordering apical 
and lateral areas; stramineous, laevigate spot at each basal angle 
diifuse or wanting. More slender species; width of body slightly 
or considerably less than one-half length ,.... 5 

2. Head subequal in length to pronotum. First antennal segment not 

extending beyond apex of head. Lsevigate spot on postfrenal 
projection of scutellum generally impunctate; Isevigate spots at 
basal angles usually as large as apical spot. An obsolescent sulcus 
barely extending to caudal margin of third abdominal stemite. 

, generous St41 

Head markedly shorter than pronotum. First antennal segment 
extending by about one-third its length beyond apex of head, 
Lsevigate spot on postfrenal part of scutellum conspicuously 
punctured; Isevigate spots at basal angles much smaller than apical. 
A well-defined median sulcus extending to apex of fourth or to 
about middle of fifth abdominal stemite 3 

78» 3 Uichanco: A revision of the genm Ectenus 287 

3. Lateral angles of terminal abdominal segments only moderately 

dentate, those of sixth armed with a short spine. External 
genitalia of female broadly rounded at apices. Lateral halves of 

propleura very deeply and thickly punctate 

mesoleucus Bergi'oth 

Lateral angles of terminal abdominal segments very prominently 
dentate and armed with rather long spines. External genitalia of 
female acute. Propleura very sparsely and shallowly punctate, 
except for a small irregular patch near lateral margin 4 

4. Second antennal segment slightly shorter than basal pudicus Stal 

Second antennal segment about one-half longer than basal 

spinosus sp. nov. 

5. Anterior angle of pronotum with a conspicuous ectally directed 

denticle; humeral angle more prominent, subacute; anterior area of 
pronotum broadly subconfluent with posterior. Second antennal 

segment somewhat longer than third 6 

Anterior angle of pronotum without prominent denticle; humeral angle 
of pronotum less salient, somewhat rounded; anterior area of 
pronotum more strongly gibbous, between lateral one-fourth 
distinctly delimited from posterior by a transverse uniserially 
punctate impression. Second and third antennal segments usually 
subequal in length spectabilis (Burmeister) 

6. Rostrum extending to about middle of fifth abdominal sternite. 

Median sulcus extending to base of, or midway into, fifth abdominal 

sternite xreus Stal 

Rostrum extending to, or only a little beyond, caudal margin of third 
abdominal sternite. Median ventral sulcus on abdomen shorter 
or posteriorly more obsolescent brevirostris Stal 


Stal. Ann. Soc. Ent. France, Ser. 4 (1865) 167; 
Ofv, Vetensk.-Akad. Forhandlingar (1870) 622. 

This species is apparently quite rare. There are available 
to me only three specimens. All of these were collected from 
the coral islands of the Sulu Archipelago, wherein the species is 
probably endemic. 

JOLO ISLAND, Bud Dajo (A. Duyag), 1 ^ (A. Lopez), 9 ; Bana- 
RAN Island, Tawi-Tawi Group {A. Duyag), 1 S. 


Bergroth, Ann. Soc. Ent. Belgique 57 (1913) 150-151. 
Closely resembling E. generosus in size, body conformation, 
and general color pattern, but the two species are readily 
separable with the aid of the key. In addition, the male anal 
segment is deeply sinuate in generosus, while in mesoleucus it 
is only broadly so. 

288 The Philippine Journal of Science ^^* 

POLILLO, October, 1909, B. S. Ace. No. 13031 {McGregor), 1 
^, 1 $ (paratypes) ; Luzon, Mt. Maquiling, elevation 350 
meters, September 23, 1932 (P. Obien ), 1 $ ; elevation 
900 meters, September 4, 1932 (/. Alhama), 1 $ ; Mt. Ba- 
nahao, March 23, 1907 (Banks), 1 9 ; 1914 (/. Valdez), 
2 ? ; Nueva Vizcaya (Alzapan and Ramos), 1 S. 

Three males and 4 females (one female bearing identification 
label in Dean C. F. Baker's handwriting), collected from Los 
Baiios, Laguna, by various collectors, May, September and Oc- 
tober, 1914 to 1925. Up to about the latter year, "Los Baiios" 
was used in a broad sense in the insect labels of the College of 
Agriculture collection, and was often made to include material 
from Mt. Maquiling at different elevations. 


Length of body 17 millimeters, width between humeral angles 
7. Ground color above dark green, with light brownish oliva- 
ceous. Other characters as given in key above. 

Closest apparently to Ectenus pudicus Stal from Mysol and 
Waigiu, but differing from this principally by the second an- 
tennal segment being considerably longer than basal, in color 
characters and in the more densely and deeply punctate body 

Negros Island, Negros Oriental, near Lake Dako (collector?), 
in the Bureau of Science collection, 1 ? (type, burnt in 1945). 


Burmeister, Nova Acta Acad. Leopold. IS, Suppl. 1 (1834) 291, PL 
51, fig. 5 {Cimex) ; Dallas, List of the specimens of hemipterous in- 
sects 1 (1851) 174, PI. 5, fig. 5; Stal, Ofversigt Vetensk.-Akad. 
Fordhandlingar (1870) 621. 

Two specimens in the lot before me have antennal segment 
2 somewhat longer than 3 and a third specimen has the humeral 
angles subacute, as in s^reits Stal; but the sum of the other 
characteristics makes their identity as spectabilis beyond 

Luzon, Laguna, Mt. Maquiling, mainly at and near summit, 
1070 meters, occasionally at lower elevations, 13 ^ ^ , 40 ? ^ 
(various student collectors), mostly during rainy season, from 
June to October; Tayabas, Quezon National Park, near Atimo- 
nan, April, 1931 (Jose Campo), 3 ^ ^, 5 ?9; Mt Bulusan, 
elev. 350-800 m. (F. A. Maclang), 2 ^^. Apparently endemic 
in southern Luzon, where it is fairly common on Mounts Ma- 
quiling and Banahao. 


Stal, Ofversigt Kongl. Vetenskaps.-Akad. Forhandlingar (1870) 621. 

78»3 Uichanco: A revision of the genus Ectenus 289 

BOHOL, May 1906 {A. Celestino) 9. Card says "Cerambyci- 
dae". Mindanao, Agusan River, Dec. 27, 1909 {A. Celes- 
tino), 1 9f October 1, 1910 (Schultze), 9 $; Davao; 
November, 1911 (C. M. Weber, 1 $ ; September, 1911 
(C. M. Weber) ; $ Biliran, North of Leyte, Naval, 1914 
(McGregor), 1 $. 


Stal Ofversigt. Kongl. Vetenskaps. Akad. Forh. (1870) 622. 

This species was not included in the collection before me. 
It is known only from St^Fs type specimen, which, according 
to Prof. Dr. 0. Lundblad (in lit, October 18, 1948), is extant 
in the Royal Natural History Museum of Stockholm. Accord- 
ing to the same informant, the type locality is Manila. 
"Manila" in StaFs time was loosely used by European authors 
to denote the Philippines; not necesarily any definite collecting 
locality. No authentic capture of any species of Ectenus has 
ever been made in Manila. 

Because my pertinent references were destroyed in 1945, I 
have been unable to recheck the literature on this species. I 
am, therefore, indebted to my friend. Prof. Robert L. Usinger, 
the well-known hemipterologist of the University of California 
at Berkeley, who very kindly transmitted to me from London 
(September 14, 1948), the following helpful information: 

"Curiously enough, brevirostris Stal is not mentioned, even in synonymy, 
with other species treated by Stal in Enumeratio Hemipterorum, part V, p. 
43. This is quite surprising, since he mentions sereus which was 
described in the same publication [supra] on a previous page in 1870. . . 
Unfortunately, the species is not represented in the British Museum 
collection. . . . 

"Stil does not number brevirostris— but treats it after *2. E. sereus 
St&l. . . .' Following the description of wreus he says, 'Specimen unum 
divergit rostro breviore, apicem segmenti tertii ventris baud vel pauUo 
superante, articulo tertio quarto dimidio longiore, sulco ventris breviore 
vel posterius obsoletiore. An species distincta, E. brevirostris StaF " 

Since the completion of the present manuscript, a set of Stales 
Enumeratio Hemipterorum and also Stal's Hemiptera insuUirum 
philippinarum were received in the library of the College of 
Agriculture through the courtesy of the Kungl. Vetenskapsaka- 
demiens Bibliotek, Stockholm. 


By Dhirendra Nath Chakraverti 

Botany Department, R. G. Kar Medical College, Calcutta 

(Formerly Carmichael Medical College, Calcutta) 


In the present paper the writer records the result of detailed 
investigation of anomalous secondary growth in some roots of 
Hydrocotyle asiatica L. (the Asiatic Pennywort) of the family 
Umbelliferae, a preliminary account of which was published 
some time back.^ The anomaly consists in the fission of the 
vascular cylinder into numerous discrete vascular bundles at 
the base of some roots of this plant This stoloniferous herb 
is common in Bengal and is found in all parts of India at 
altitudes up to 2,000 feet, its distribution being tropical and 
subtropical (Hooker) J 

Naked eye observations of the roots do not, as a rule, show 
any indication of the fission of the vascular cylinder inside the 
same but in some cases the cleavage may be inferred from a 
slight wavy outline or a little flattening of the root itself. 


For microtome sections the selected roots were cut into bits 
from the base to the tip and fixed in three lots: the first in 
stock chromo-acetic solution, the second in *CrAF,' named by 
Randolph ^^ and the third in formalin-water (40 per cent for- 
malin 10 cc and distilled water 90 cc) at GO"" C. for 2 hours. 
Specimens killed and fixed in the first fixative were treated in 
the usual way; those of the second lot were directly immersed in 
75 per cent alcohol, then passed through the alcohol grades, 
dehydrated, cleared in cedar-wood oil and embedded as usual; 
and the last lot, fixed in formalin-water, was treated directly 
with 95 per cent alcohol for 1 kour, then passed through pure 
dioxan, 2 changes of 1 hour each, dioxan-paraffin (1: 1) 1 hour, 
paraffin 2 changes of 15 to 20 minutes each, and finally embedded 
according to Chapman^s ^ review of T, Garvin. All the fixatives 
proved equally successful. Serial microtome sections were cut 

9543 2 


292 The Philippine Journal of Science *»** 

in transverse plane 8 to /n to 19 /* thick from the base of the root 
towards its apical part and some longitudinal sections were also 
made for comparison. Delafield's haematoxylin and (rapid) 
saf ranin as suggested by Dean, * safranin and light green dis- 
solved in clove oil and Bismarch brown and light green were 
used. The first combination gave the best result. 

For the microscopical examination of the cleft xylem-mass in 
situ, a few roots were decolorised after Debenham ^ in 60 per cent 
lactic acid for some days, washed in 75 per cent glycerine for 2 
to S days, then treated with 50 per cent alcohol to ensure complete 
washing out of the acid and glycerine used. The materials were 
afterwards bleached in Eau-de-javelle at 40"" C. for about 15 to 
20 minutes followed by a thorough washing in water for 12 hours 
with several changes and then treated with 10 per cent alcohol ; 
the roots were then stained as a whole without any sectioning. 
The writer, however, got equally good results by gently boiling 
the materials in 60 per cent to 80 per cent lactic acid solution 
over water-bath for 2 to 4 hours according to the thickness of 
the roots. After the first hour's boiling some more lactic acid 
solution was added at intervals to maintain the uniform strength 
of the boiling solution. This hastened the preliminary decolor- 
ization without any perceptible injury of the materials and was 
a time-saving modification, at least in these materials. Free- 
hand sections from the base, middle and apical regions of many 
of these roots were also examined mostly mounted in dactic acid. 


The fission of the xylem-mass is found confined to the base 
of some roots of 1 to 3 mm in diameter, within a space of about 
3 cm from the point of their origin; beyond this the nature 
and arrangement of the xylem-masses are normal up to the tip. 
The fission occurs, generally, only once in the root but very 
rarely, a second fission is also noticed in the same root. On 
examining the serial microtome sections from the neighborhood 
of the tip region the apical region shows radial arrangement of 
the vascular bundles, which may be three, four (PI. 2, fig. 1) or 
five in number in different specimens. The pith is distinct, 
parenchymatous and thin-walled. In the apical region of some 
young roots the pericycle has been found to be 2 to 3 layers thick 
(PI. 2, fig. 1) but in other older roots the pericycle has become 
3 to 4 layers in thickness. The major portion of the root shows 

^*' ^ Chakraverti: Roots of Hydrocotyle asiatica Linn, £93 

secondary growth and thickening of the dicotyledonous type. 
In adult roots, in many cases, the center is occupied by meta- 
xylem elements (PL 2, fig. 2). 

In the middle portion of the root, the cambium approximates 
a circular ring which towards the basal region becomes pro- 
gressively invaginated towards the center between the consecu- 
tive xylem bundles (PL 2, figs. 2, 3, 3a and PL 1, fig. 2). 
Ultimately, it breaks up into as many separate arcs as there are 
primary xylem bundles. Each of these separate cambial arcs 
then extends round and encloses the primary xylem with some 
amount of secondary wood in an annular manner (PL 2, figs. 
4 and 4a). The smaller ring of cambium, thus formed, then 
produces at its sides and towards its center, a large amount 
of parenchymatous cells, arranged in regularly widening circles 
which push apart the original xylem-masses laterally from one 
another and force them away from the center, thereby causing 
the cleavage or fission of the original xylem-mass (PL 1, figs. 1 
and 2). In the absence of such cambial growth the xylem- 
masses would have formed a continuous cylinder. This fission 
causes a slight flattening or wavy outline of these roots as 
already mentioned earlier. The xylem parenchyma is found 
abundantly between the cleft xylem-masses and also in the 
secondary wood. 

The old roots in general show copious development of paren- 
chyma. The primary medullary rays at the back of the proto- 
xylem widen gradually towards the cortex to form somewhat 
funnel-shaped medullary rays (PL 1, figs. 1 and 2). The pri- 
mary phloem bundles, composed of thick-walled cells, are situated 
at the outer end of the broad radially elongated medullary rays 
alternating with the primary xylem bundles (PL 1, fig. 2). The 
secondary phloem bundles are collateral, often truncated, conical 
in shape and are separated from one another by the widening 
medullary rays. Sometimes, the walls of these secondary 
phloem bundles are thickened but they give cellulose reaction. 
Starch grains are found deposited in abundance in all the paren- 
chymatous elements within the stele (PL 1, fig. 2); calcium 
oxalate occurs very rarely as sphaeraphides in some parenchy- 
matous cells (PL 2, fig. 5.) 

The cortex is gradually exfoliated from the old roots (PL 1, 
fig. 2), but in the younger ones the cortical cells are found 
very loosely arranged with broad air-spaces (PL 1, fig. 1). 
Phellogen appears in the pericycle after secondary growth (PL 1, 

294 The Philippine Journal of Science i^*^ 

%• 2, and PL 2, fig. 5) and the continuity of the endodermal 
layer is interrupted, its cells becoming flattened in the mean- 
time (PI. 1, fig. 5). Glandular ducts are found opposite the 
xylem and the phloem groups of the primary vascular strands 
(PL 2, fig. 1) as noted by Solereder.^* After the secondary 
growth such ducts are also seen external to the secondary 
phloem (PL 1, figs. 1 and 2; and PL 2, fig. 5). 

A somewhat similar fission of the xylem-mass was reported 
by Koch ^^ in the mature napiform roots of Sedum maximum 
Sut. of the family Crassulaceae and its allies, the difference 
being that in this case the fission occured in the middle portion 
instead of the basal region of our roots and the bundles were 
hadro-centric instead of the collateral ones as are found in the 
roots of Hydrocotyle asiatica L. 

The fission of the xylem-mass in the root of Hydrocotyle 
asiatica L. differs in many ways from that of Azorella selago 
Hook. f. as noted by Ternetz,^^ although the two genera belong 
to the same sub-family Hydrocotyloideae under the family 
Umbelliferae. In Azorella disorganization of the ground tis- 
sues of the secondary cortex takes place and is followed by the 
cleavage of the xylem-mass. Here a meristematic tissue origi- 
nates secondarily in the parenchyma bordering the cleft places 
whereas in the roots of Hydrocotyle, disorganization of the 
parenchsntnatous ground tissue has not been observed nor any 
new meristem has been found to develop anywhere. In Hydro- 
cotyle the conducting cylinder of the root splits up into as many 
segments as there are primary xylem bundles and all these 
separate strands, thus formed, are found arranged in a regular 
ring unlike the innumerable, irregularly scattered vascular 
strands of Azorella. 

The anomalous root structure of Oenanthe, Magydaris and 
Myrrhis of the same family (Umbelliferae) is, however, 
of a quite different nature (Solereder).^* Fission of the 
xylem-mass in roots of plants belonging to other families has 
also been reported from time to time. Dastur and Kapadia^ 
have observed the fissoin of xylem caused by the dilatation 
of parenchyma in the aerial and the underground roots of Tinos- 
pora cordifolia Miers. (Family Menisperm^iceae) . Recently, 
P. €. Joshi ^® has noted the splitting up of the stele of the old 
stems and roots of Thylacospermum rupifragum Schrenk., a 
Tibetan Caryophyllaceous plant, into numerous irregular vas- 
cular strands. A. C. Joshi ^ has reported a somewhat similar 
anomaly in the roots of another Tibetan plant, Stellera chamae- 

'*» 5 Chakraverti: Roots of Hydrocotyle asiatica Linn. 295 

jasme L. of the family Thymelaeaceae. Solereder^^ records 
the occurrence of the cleft xylem-mass in the stems and in some 
cases in the roots also of some plants belonging to the families 
Caryophyllaceae, Malpighiaceae, Sapindaceae, Caesalpinieae 
{Bauhinia)f Umbelliferae, Asclepiadeae, Convolvulaceae, Big- 
noniaceae, Acanthaceae and Aristolochiaceae. 


In great many cases secondary differentiation of the stelar 
tissues of the dicotyledonous roots has been found to deviate 
more or less considerably from that of the normal type. All 
stages, from the simpler cases of the unequal growth of the 
xylem through its furrowed appearance to the splitting up of 
the vascular cylinder into separate strands have been observed 
in a number of families by different authors. The segments, 
so formed, may be arranged simply in a regular ring as in the 
roots of Hydrocotyle, or by repeated cleavage and secondary 
growth they may produce masses of wood and bast irregularly 
interwoven with one another as in Azorella. Such anomalous 
vascular structures have also been observed in the stems of some 
tropical lianes belonging mostly to the families of Combretaceae, 
Malpighiaceae and Bauhineae (De Bary).^ Another type of 
cleavage has been noted by Lashevsky^^ in the underground 
stems of Daphne julia (Family Thymelaeaceae) where the 
fission is caused by the activity of the parenchyma of the 
pith, wood and wood-rays. The cause of these anomalies has, 
however, not been satisfactorily explained. Lashevsky thinks 
that this is an ancestral character retained by the species show- 
ing the anomaly but in the present case, as in all others, where 
great parenchymatous development with abundance of starch 
grains as the cell-contents is the consequence of such cleavage, 
the explanation put forward by Haberlandf^ appears to be 
very reasonably. He maintains that such peculiarities of sec- 
ondary growth serve to ensure the production of the storage 
tissue characteristic of the roots of many Umbelliferae, render- 
ing the deposition and renewal of reserve materials easier and 
more effective after the intermixing of the conducting and the 
storage tissues. 


1. The roots of Hydrocotyle asiatica L. show usually normal 
secondary growth and thickening characteristic of the dicoty- 
ledonous root with a great tendency towards the development of 
the parenchymatous tissues. 

296 The Philippine Journal of Science ^^^ 

2* In some roots, however, the vascual cylinder at the basal 
region split up into numerous discrete vascular strands, their 
number being the same as the number of the primary vascular 
bundles of the root. Here the continuous cambial ring becomes 
progressively invaginated towards the center of the root and 
breaks up into separate arcs, each one of which ultimately 
encloses a xylem-mass including one primary xylem bundle. 

3. The fission of the vascular cylinder is probably due to the 
tendency of the development of storage parenchyma, thus ren- 
dering the deposition and renewal of food materials easier and 
more effective in the roots. 

In conclusion, I desire to express my grateful thanks to Dr. S. 
R. Bose, Professor of Botany of this College, for his valuable 
help and criticisms in this investigation. 


1. Chapman, G. H. The use of dioxan in histological technique. Stain 

Tech. 14 (1939) 113. 

2. Chakraverti, D. N. A preliminary note on the fission of vascular 

cylinder in some of the roots of Hydrocotyle asiatica Linn. Curr. 
Sci. 9 (1940) 230-233. 

3. Dastub, R. H. and G. A. Kapadia. The anatomy of climbing plants, 

Jour. Ind. Bot. Soc. 10 (1931) 110-121. 

4. Dean, H. L. Delafield's haematoxylin and safranin for staining 

plant materials. Stain Tech. 15 (1940) 61-65. 

5. Be Bary, a. Comparative Anatomy of the Phanerogams and Ferns. 

Oxford. (1884). 

6. Debenham, E. M. a modified technique for the microscopic examina- 

tion of the xylem of whole plants or plant organs. Ann. Bot. N. S. 
3 (1939) 369-373. 
7» Habbrlandt, G. Physiological Plant Anatomy. London. (1914). 

8. Hooker, Sir J. D. Flora of British India 2 (1879) 669. 

9. JOSHI, A. C. Secondary thickening in the stem and root of Stellera 

chamaejasmae L. Proc. Ind. Acad. Sci. 2 (1935) 424-436. 

10. JosHi, P. €. Anatomy of the vegetative parts of two Tibetan Caryo* 

phyllaceae-^Arenaria musdformis Wall, and Thylacospermum rupi" 
fragum Schrenk. Proc. Ind. Acad. Sci. 4 (1936) 52-65. 

11. KoGH, L. Entwickl. d. Crass. Verh. naturhist.-med. Ver. zu Heidel- 

berg. Bd. 1, 4 Kelt. (1876). 

12. Lashevsky, V. On the liane structure in the subterranean stem of 

Daphne julia. Bot. Abstracts, April-May (1926). 

13. Randolph, L. F. A new fixing fluid and a revised schedule for the 

paraffin method in plant cytology. Stain Tech 10 (1935) 95. 

14. SouiREDEp, H. Systematic Anatomy of the Dicotyledons. Oxford 


15. TiaiNETZ, Ch. Morphol. u. Anat. d. Azorella selago Hook, f, Bot. Zeit. 

60 (1902) 1-20 und Tab. 1. 


(Explanation of the abbreviation of the parts labelled in the figures is 
given at the end. Figures of Plate 1 are photomicrographs and those of 
Plate 2, are camera lucida drawings from the roots of Hydrocotyle asiatica 

Plate 1 

Fig. 1. Transverse section of a root showing earlier stage of the formation 
of cambium ring around each xylem-mass including one primary 
xylem. A cork-cambium is forming in the pericycle. The cortex 
is in process of sloughing off especially in the lower side. (Eye 
piece 7.5 X and objective 16 mm.) 
2. Transverse section from the base of an old root showing four dis- 
crete xylem-masses arranged in a ring. Pericycle 3 to 4 layered 
showing phellogen outside. Endodermis interrupted at places. 
Cortex and epidermis completely sloughed off. (Eye-piece 5 X 
and objective 16 mm.) 

Plate 2 

Fig. 1. Transverse section from the apical region of a young root showing 
tetrarch arrangement of vascular bundles and 1 to 3 layers of 
pericycle. X 430. 

2. A portion of the transverse section from the middle region of an 

adult root after secondary growth showing the invaginating 
cambium and the metaxylem elements meeting at the center. 
X 450. 

3. Transverse section from the basal part of an old root giving semi- 

diagrammatic view of the normal circular cambium becoming 
progressively invaginated towards the center. Also it shows 
among other parts a space within for 3 to 4 layered pericycle 
the inner limit of which has been shown in a circle of dots. 
X 80. 
3a. One of the three xylem bundles of Fig. 3 showing details of a 
portion within the invaginated cambium and a portion of the 
primary medullary ray outside. X 430. 

4. Semi-diagrammatic view of the root in transverse section showing 

the formation of a complete ring of cambium round each xylem- 
mass. The continuity of the endodermis is found interrupted 
at several places of the section. Just below this phellogen is 
shown in circular outline in dots. The innermost three small 
circles in dots represent the outer limit of the discrete vascular 
bundles. The bigger circle in dots enclosing the three separate 
vascular bundles is the inner limit of pericycle which is 3 to 4 
layers of cells in thickness. X 70. 
4a. One of the three xylem-masses of Fig. 4, shown in details. Here 
a few cells of the basal part of the primary medullary rays 
have been shown just opposite to protoxylem. X 440. 

5. Drawing of an outer part of PI. 1, fig. 2, showing interrupted 

endodermis, phellogen, glandular ducts, sphaeraphides, etc. X 



The Philippine Journal of Science 



a — air-space 

c — cortex 

cam— -cambium 

dvb — discrete vascular bundle 

e — epidermis 

en — endodermis 

gd — ^glandular duct 

mr — medullary rays 

mx — ^metaxylem 

p — pith cells 

pc — ^pericycle 

ph — phloem 

phi — phellogen 

pph — ^primary phloem 

ptx — ^protoxylem 

px — primary xylem 

sg — starch grains in parenchyma- 
tous cells 

s ph — ^secondary phloem elements 

sx — secondary xylem elements 

X — both the primary and secondary 
xylem elements 

xp — ^xylem parenchyma 

Chakraverti: Roots of Hydrocotyle Asiatica Linn.] [Philip. Jour. Sci., 78, No. 3. 






Chakraverti: Roots of Hydrocotyle Asiatica Linn.] [Philip. Jour. Sci., 78, No. 3. 




Director of the Unilever Research Laboratorium 
Zwijndrechty Holland 

During the "Journees scientifiques des corps gras alimen- 
taires" held in April 1949 in Paris Dr. Autret of the "Centre 
National des Etudes et Recherches sur la Nutrition et FAlimen- 
tation" recorded the extremely high content of carotene in the 
fruits of Momordica cochinchinensis Spreng, called by the An- 
namites "day-gac". According to the minutes of the meeting 
in Oleagineux, 4, (1949) 309, 8 grams of carotene could be found 
in 1 kg (0.8 per cent). Further details will be published by 
Autret in the same journal. 

As we are keenly interested in materials with a high carotene 
content, the investigation of this remarkable fruit was of great 
value and we succeeded in getting this fruit— carefully sealed in 
beeswax — ^from the Philippines.* 

The fruit resembles in shape a great pear and in color an 
orange. The weight was about 300 grams. On first sight our 
attention was drawn by the deep red layer surrounding each of 
the 14 black seeds within the fruit pulp. According to the 
Carr-Price reaction this layer contains carotenoids, but analysis 
showed that carotene itself was present in only a small quantity. 

The fruit was divided in four parts and the total carotene 
content of each determined: 

a. Skin 

b. Fruit pulp (pale yellow) 

c. Pulp round seeds (blood red) 

d. Seeds (black) 



46 g. 

20 g. 
31 g. 

295 g. 

P«r cent 
of total 







1.62 mg. 
0.82 mg. 
0.83 mg. 

3.27 mg. 

Per cent 





* We are very grateful to Dr. Eduardo Quisumbing, Director of the 
National Museum in Manila, Mr. Kenneth B. Day, Director, and Mr. 
G. J. Wouters, Asst. Technical Director of the Philippine Refining Co,, 
Manila, for all the trouble taken in sending this fruit to Europe in a fresh 


300 The Philippine Jommal of Science ^^^ 

The results are nearly negative, and we see once more that 
people are often attracted by the deep red color of some fruits, 
but that this is not always the sign of a high content of provi- 
tamin A in the form of carotene. 

From these results we may conclude that Autret's statement 
must be based on an error, probably caused by the deep red 
color of the pulp. The coloring matter was not further in- 
vestigated because the carotene content showed to be in no way 





lloilo College, University of the Philippines^ Iloilo City 



The genital systems of the four groups, namely, Scylla serrata 
(Forskal) ; S, serrata var. paramamosain Estampador; S. ocean- 
ica (Dana) and S. tranquebarica (Fabricius), do not show 
marked diflferences in their gross morphological features. 
Hence, the following general description will serve for all. 

The abdomen of the male (Plate 2, fig. 1) is much narrower 
than that of the female, and is provided with only two pairs of 
appendages which are modified as copulatory organs. The ex- 
ternal genitalia consist of a pair of tubules or ducts, which are 
about eight millimeters long in a crab that measures nine centi- 
meters across the widest portion of the carapace. Each duct 
comes out from an opening at the base of the thoracic leg, and 
is received into the troughlike excavated area (dsterna sper- 
matica) on the inner surface of the base of the terminal joint 
of the first abdominal appendage. 

The first abdominal appendage (Plate 2, figs. 2, 3) is composed 
of two joints; the basal one is broad, rectangular, and flattened 
on its inner surface, which is held appressed to the sternal 
wall. The terminal segment has a sinuous outline; it is long 
and tapering with the point shaped like that of a quill pen or 
that of a hypodermic needle. The cisterna spermatica (Plate 
2, fig. 2 c) apparently functions as a receptacle to receive the 
sexual products expelled from the ejaculatory duct. From this 
cistern a canal leads ventrally, that is, towards the outer surface 
and to the mouth of the funnel-tube — ^the remaining portion of 
the last segment, which is so constructed as to present a sem- 
blance to a slender funnel Indeed, its structural peculiarities 
and its relations with the ejaculatory duct point to its probable 
function of transferring spermatozoa into the vulva of the 
female in copulation. 

The second abdominal appendage (Plate 2, fig. 4) is a delicate, 
unsegmented structure. In the same animal, it measures thir- 

♦ Pt. I. Revision of the Genus, Philip. Jour. Sci., 78 No. 1 (1949) 95-109. 


302 The Philippine Journal of Science i^*^ 

teen millimeters in length, and hardly one millimeter in diameter 
at the base of the fork. One branch of the fork is spooned and 
provided with spinules; the outer branch (c), which contains a 
troughlike cavity, is inserted into the opening of the funnel- 
tube of the first appendage. 

During copulation, the ejaculatory duct pours the male sexual 
products into the cisterna spermatica which is connected to the 
opening of the funnel-tube by a canal. The forked rodlike 
second abdominal appendage presumably helps by keeping the 
spermatophores confined in the funnel-tube, in forcing the male 
sexual products into the vulva of the female. 

The testes (Plate 2, fig. 6) are paired and symmetrical. The 
two halves are connected immediately behind the foregut in 
almost the same situation as the ovaries in the female. They 
are symmetrical in position and lie inmiediately beneath the 
hypodermis. The size and shape vary with the condition of 
sexual maturity of the crab. They are small in immature speci- 
mens; while in sexually mature individuals, during the spawning 
season, they are massive, practically filling up the angular space 
between the carapace and the sternal wall, and may cover up 
almost the whole of the digestive gland. Each testis describes 
an arc following the outline of the carapace ; the terminal portion 
extending as far as the postero-lateral teeth. 

At the inner extremity, at about the region of the cardiac 
foregut, it dips inward and behind the pyloric foregut, it forms 
a prolongation that joins a similar portion from the other 
testis forming a sort of a bridge. Posterior to this bridge, it 
connects with a very much coiled tube, the vas deferens. At 
the point just above the level of the pyloric foregut, the duct 
turns inward and follows the course along the digestive tract 
It becomes narrower and continues backward to the base of 
the last thoracic leg, where it empties into the exterior through 
an opening on the coxopodite of the last leg. We shall call 
this portion the ejaculatory duct. This is protected by the 
extension of the arthrodial membrane from the base of the leg, 
the membrane rolling around it at the terminal portion. The 
duct lies closely appressed to the sternal wall and is covered by 
the lateral extension of the third segment of the abdomen. Its 
tip is received into the opening of the funnel-tube of the first 
abdominal appendage. 

There are several factors which presumably lead to the expul- 
sion of the s^ual products into the funnels tube. Undoubtedly, 

^«» 3 Estampador: Studies on Spermatogenesis and Oogenesis 303 

gravity plays an important role. So does pressui'e; this may 
arise from two sources, namely, internal and external. The first, 
we may qualify as the transmitted pressure which operates in the 
same manner as in a water-main. Pressure from outside is 
brought about by the movements of the copulatory apparatus 
during mating. In these movements, the ejaculatory duct is 
caught in between the sternal wall and the lateral extension of 
the wall, the cisterna spermatica. Other factors may be sought 
in the structural constitution of the duct itself. The epithelial 
cells that line the cavity of the ejaculatory duct are glandular 
and ciliated (Plate 2, fig. 7). Obviously, they produce some 
kind of secretion, the nature of which is not known at present. 
The cilia, of course, help in the movement of some metabolic 
products in the duct. 

As in the ovary, the testicular follicles are closely packed 
together. The lumen of each follicle is continuous with that 
of the vas deferens. The cells that line the lumen are of three 
types, namely, the spermatoblast, the nurse cells, and the secre- 
tory cells. The last are probably collagen-forming cells. 

The spermatoblasts are easily distinguished by the large 
amount of cytoplasmic materials present and by their large 
nuclei. The nurse cells are smaller, with distinctly smaller 
nuclei, and are more or less vacuolated. In both of these kinds 
of cells, the nuclei are basal in position. 

The secretory cell is big, rounded in outline and possesses 
a big vesicle. Its metaplasm takes up a bluish tint with iron 
hsematoxylin, acid fuchsin and anilin blue stain combination; 
the secretory product is apparently collagen or a related sub- 
stance. The secretory cells present in the follicles are very much 
fewer and are smaller, compared to those present in the ovarian 
follicles (Plate 7, fig. 9). 

The secretion presumably furnishes the necessary medium for 
the spermatozoa on their way to the outside. If such be the 
case, this medium must possess a certain degree of viscosity to 
protect the spermatozoa from the action of the water. In the 
case of the female, the secretory cells furnish the sticky mate- 
rial by means of which the eggs are attached to the endopodite 
setse. Considering the large number of eggs and their relatively 
much bigger size, a considerably much greater amount of ma- 
terial is needed. Perhaps this fact may explain the presence 
in greater number of secretory cells in the ovarian follicles. 

304 The Philippine Journal of Science i^' 


The spermatoblasts (auxocytes) may proceed in their develop- 
ment either singly or in groups. In either case, the growing 
auxocyte draws some of the cytoplasmic and nuclear materials 
from the surrounding cells and uses them for its growth. In 
the case where they are in groups, the competition among them- 
selves results in some being used up by the more vigorous 
auxocytes. The group is held together by a sort of cyst, prob- 
ably formed from the detritus from the used-up cells, and 
possibly cemented together by the secretion of the secretory cells. 
Such a group of auxocytes may be termed spermatocyst (Plate 
5, fig. 1 and Plate 6, fig. 6) • This group formation is frequently 
met with in the mamosain and paramamosain groups, and 
occasionally in tiie parabanhawin {S. tranquebarica} . It does 
not seem to occur in the true banhawin (S. oceanica) . 

The most distinctive changes that take place during the growth 
of the auxocyte can be observed in the increase of its bulk due 
to the increase in volume of its cytoplasm and in the increase 
in size of the nucleus. At the termination of the growth period, 
the cell undergoes transformation in two distinct ways; one, 
the whole cell, or the greater portion of it, participates actively; 
the other, the apparent activity is seemingly centered around the 
nucleus and the rest of the cell undergoes no radical structural 
changes, except for the sloughing off of the csrtoplasm. 

Heteropycnosis. — ^A very peculiar characteristic is seen in 
what is termed "heteropycnosis,*' in which the sex-chromosomes 
become greatly condensed and form an intensely staining mass, 
appearing as a karyosome, or "chromosome-nucleolus," in sharp 
contrast to the threadlike or diffused and slightly staining 
ordinary chromosomes. It appears to be a very conspicuous 
feature during the growth period and, in some cases, it appears 
even as late as in some phases during spermatid-transformation 
(Plate 5, figs. 4, 5). 

This condensed mass of chromatin materials give rise to the 
•*hetero-chromosomes*' of some authors. Among the groups of 
crabs under consideration, the chromosome-nucleolus represents 
either the XX- or the XY-pair of chromosomes* 

Spermiogenesis. — ^After the growth period, the cell undergoes 
a series of transformations; the subsequent developmental phase 
may be termed **spermiogenesis'' or "spermioteleosis.'' In S. 
tranquebarica (parabanhawin) and in the variety paramamo^ 
sain, the process of spermatogenesis can be subdivided quite 

^*' 3 Estampador: Studies on Spermatogenesis and Oogenesis 305 

distinctly into four general phases, namely, (1) the auxocyte, 
characterized by growth, (2) the pro-spermatid stage, charac- 
terized by reorganization, differentiation, and sloughing off 
of some of the materials leading to the formation of the 
definitive spermatid body, (3) spermatid transformation, also 
featured by reorganization, differentiation and further sloughing 
off of the materials, and (4) the spermatozoon. While in S. 
oceanica and S. serrata the four general phases cannot be dis- 
tinctly differentiated, there is a considerable merging of these 
phases; hence, an apparent shortening of the process of sperma- 


This group is described first, because the two types of sper- 
miogenesis previously referred to occur in a somewhat more 
elaborate manner. The two types are linked with the chromo- 
somal constitution of the germ cells involved: (1) in one class, 
there is tetrad formation and almost the whole of the cell ac- 
tively participates; (2) in sharp contrast to the other class, 
the apparent activity is seemingly centered in the nucleus. We 
shall refer to the first as having the tetrad formation and to 
the second as without tetrad formation. 

I. Spermiogenesis in cells having tetrad formation. — In this 
class the process involves telokinetic movements which include: 
(1) movements of th6 nucleus, (2) extrusion of some of the 
nuclear materials and most of the cytoplasm, (3) peculiar be- 
haviour and movements of the acroblast, (4) rotation of the 
chromosomes. This phenomenon is especially noticeable in the 

Some of the more obvious changes that take place in the 
series of transformations leading to the formation of sperma- 
tozoa may be seen in the following illustrations : 

Plate 3, fig. 1, represents a growing auxocyte. Within its 
body, the shriveled nucleus of a nurse cell can still be seen. Its 
nucleus is apparently in the leptotene-stage, the spireme threads 
beginning to arrange side by side, evidently in preparation for 
synapsis (syndesis). The tetrad formation is quite obvious; 
this is shown by a loop giving rise to four chromatids. It aeema 
that in this case synizesis is foreshadowed by a migration of 
the prochromosomes towards certain places along the periphery 
of the nuclear wall, where they become closely aggregated 
before the leptotene threads are spun out from them. The big- 
gest of these pre-sinizetic knots apparently give rise to the 

306 The Philippine Journal of Science i^« 

Plate 3, fig. 2, shows the surface view of an auxocyte at the 
termination of the growth period. Evidently, it is in the con- 
traction phase; the chromosome-nucleolus is at the center, and 
from it, the prochromosomal threads radiate, apparently in 

Plate 3, fig. 3, depicts a pro-spermatid body, wherein the 
process of concentration is going on, apparently in preparation 
for the subsequent reorganization. From this point on, the 
subsequent changes that follow are not those of growth, as 
these are characterized by diminution of the size of the cell 
accompanied by reorganization* and differentiation. The term 
spermatocjrte does not seem to fit in this particular case, taking 
into consideration the previous history, and the changes that 
are to follow. In this figure, the spireme has been transformed 
into chromatids. The tetrad formation is indicated by the 
grouping of chromatids into four. Though in a somewhat 
diffuse condition, the arrangements in pairs is quite apparent 
as in (1) in which each mate is composed of a row of chroma- 
tids. Presumably this pair will form in synapsis an anaschistic 
rod tetrad with terminal connection. 

Plate 3, fig. 4, shows the chromosomes in pre-synaptic asso- 
ciations; the pairs are recognizable. Three types of tetrads 
are present: (1) the loop-shaped, which is apparently the X- 
chromosome in the XY-pair, the hetero-chromosomes {h); (2) 
rod-like; and (3) V-shaped. Some of the chromosomes are 
distinctly segmented. 

Plate 3, fig- 5, shows the synaptic figures; the mates of some 
differ in size, form and in structure. Hence, they are hetero- 
morphic. In the attraction sphere or idiosome (i) are darkly 
stained bodies with granular constitution; these are the pro- 
acrosome granules (p. ac). 

The changes that bring about the transformations leading to 
the formation of spermatozoa are featured by series of concen- 
tration, contraction, and diminution of both the nuclear and 
cytoplasmic materials. Some of the phases of spermatid trans- 
formation are illustrated in the following figures: 

In Plate 3, fig. 6, the pro-spermatid body has assumed a more 
definitive form, its outline has become more regular. The chro- 
matin net-knot (nucleo-chromosome) appears as a big heavily 
stained body near the center; the rest of the chromatin mate- 
rials have ranged along the border of the nuclear wall toward 
the apical pole as small deeply stained masses, while some are 

^^' ^ Estampador: Studies on Spermatogenesis and Oogenesis 307 

apparently being sloughed off at the opposite pole. The darkly 
stained mass (n. e) is presumably composed of extruded nuclear 
materials; the other {g) containing heavily stained bodies 
probably constitutes extruded Golgi-remnant bodies. The pic- 
ture shows clearly the process of diminution involving the 
cytoplasm and the karyoplasm. The attraction sphere or idio- 
some (0 is now at the apical pole and contains a heavily stained 
body, the acroblast (ac). 

Plate 3, fig. 7, depicts the process of concentration, as evid- 
enced by the broken and reticular condition of the cytosome at 
the basal portion of the pro-spermatid body. The nucleus has 
moved farther toward the apical pole; the chromosomes are 
now dyads, instead of tetrads, and the number has been reduced 
to six. The small U-shaped chromosome near the pole is 
evidently the extra sex-chromosome (m) . It behaves differently 
by moving ahead of the others. Reduction of the amount of 
the chromatin materials and, consequently, the number and 
size of the chromosomes must have taken place in the preceding 
stages. This reduction was brought about by a sort of regroup- 
ing or seggregation of the chromosome materials — ^the tetrads 
separating into dyads without undergoing division in the usual 
manner. One of the groups, evidently composed some of the 
nuclear materials that were cast off (Plate 3, fig. 6,n. e). 

Plate 3, fig. 8, shows the differentiation of the definitive 
spermatid body {sp, 6). The nuclear materials have reached 
the proximity of the attraction sphere; presumably the heavily 
stained body in the form of a nucleo-chromatin is the sex- 
chromosome (x). Apparently the acroblast has disaggregated; 
The spermatid has a form very much like that of a hat; the 
head is obviously composed of the sphere-substance referred 
to by Wilson (1924)* as a clear substance inside the idiosome; 
the brim is composed of cytoplasm, and the nucleus is apparently 
located centrally in this region. This figure indicates clearly 
that only a small portion of the cytoplasm is utilized in the 
formation of the definitive spermatid body; the rest is sloughed 
off. This mass of residual protoplasm (r. p) degenerates 
without taking any further part in the sperm formation. Some 
of these discarded materials are presumably used in the forma- 
tion of the cyst, in cases where the auxocytes are in group 
formation (spermatocyst). The remainder is released into the 
lumen of the follicle and into the spaces between the follicular 
tubules, where they are observed as the undifferentiated chro- 

9543 8 

308 The Philippine Journal of Science ^^^ 

mophyllic mass. Some sloughed-off nuclear materials and Golgi- 
remnant bodi^ are commonly seen in this mass. The heavily 
stained mass {c. e) among the cast-off materials may represent 
either nuclear materials or a portion of the cast-off acroblast. 

Plate 3, fig. 9, represents a sort of an abortive mitotic figure 
showing division involving mainly the nucleus. Part of the 
cytoplasm will undoubtedly go with the casting off of the 
daughter nucleus. Apparently, this process constitute the second 
meiosis, and reduces the chromosomes from dyads to monads. 
The small chromosome which is out of the alignment and nearer 
the acroblast, is evidently the extra sex-chromosome. It is no 
longer Unshaped (Plate 3, fig. 7, m). The acroblast {ac) is 
again aggregating. From its constitution, the cast-off daughter 
nucleus appears to be normal; whether it develops into a normal 
sperm is yet to be proved. There are indications, however, that 
it might be the case, as can be inferred from the presence within 
the remains of the auxocyte body of two spermatids or two 
spermatozoa in close proximity to each other, a condition which 
seems to be typical. 

Plate 3, fig. 10, shows the top view of a spermatid repre- 
senting a later stage than the preceding. It represents a 
figure similar to that of a conical lamp shade; the apex is 
occupied by the idiosome (i) containing two deeply stained 
bodies — ^the acroblasts, so disposed opposite each other. The 
process of concentration is going on, as indicated by the condi- 
tion of the cytoplasm. 

The subsequent concentration will result in the sloughing off 
of some materials, as shown in Plate 3, fig. 11; it also brings 
the two acroblast bodies together. These bodies do not fuse, 
however, the more homogeneous one leaning more to the outside 
of the idiosome. Further concentration and sloughing off of 
materials, apparently results in one of the acroblasts being 
pulled down towards the base to become the acroblast-remnant 
(Plate 3, fig. 12, ac. r), which eventually will be cast off. 

The other acroblast body is not shown in Plate 3, fig. 12; 
apparently it has undergone disaggregation, as indicated by the 
somewhat clear appearance of the idiosome. The figure indi- 
cates that the process of concentration is going on. This is 
subsequently followed by contraction. Plate 3, fig. 13, illustrates 
this phase, showing the basal portion viewed from below, and 
the position of the nucleus at the center of the base of the sper- 
matid body. It also shows the chromosomes which are located 
along the periphery of the nuclear wall. 

'^'3 Estampador: studies on Spermatogenesis and Oogenesis 309 

Further contraction results in the sloughing off of some more 
cytoplasmic materials. Plate 3, fig. 14, represents a stage that 
results from the preceding changes. The figure is something 
like a dome with the nucleus forming the base. Apparently the 
acroblast is in a state of disaggregation. The subsequent pro- 
cesses of concentration and contraction that follow transform 
the figure into something resembling a bowl (Plate 3, fig. 15). 
The acroblast has reaggregated and occupies a position at the 
border of the idiosome. 

In the subsequent phase, the spermatid assumes a conical form 
(Plate 3, fig, 16) and the acroblast has split into two bodies, of 
which one moved down toward the base, presumably to become 
the second acroblast-remnant. It is highly probable, however, 
that its materials may become finely disseminated and may be 
used in the formation of the nuclear cup. The other acroblast 
body remains at the apex, and it may eventually constitute the 
definitive acrosome body. 

The diminution of the cytoplasm is quite obscure from this 
point on. It seems that in these later stages, in the basal 
portion wherein the nucleus is located, the surface-tension 
balance has reached a point which would establish a certain 
degree of rigidity in this region. Consequently the sloughing 
off of materials can no longer take place in this area; it is 
shifted to the lateral surface, where the surface tension has not 
reached the determinative fixity. 

Plate 3, figs. 16 and 17, show this shifting and also that of the 
position of the definitive acrosome body towards the base. This 
apparent movement of the acrosome body is possibly brought 
about by the concentration of cytoplasmic materials which most 
likely exerts a pulling force and the subsequent contraction 
produces a further downward shift. Plate 3, fig. 18, illustrates 
the movement of the definitive acrosome. 

Plate 3, fig. 19, shows the spermatozoon; it measures 13 
microns in length and a little over 11 microns in diameter at 
its base. The form is similar to that of a funnel or a conical 
cup. The nucleus is located at the mouth of the cup; hence, 
we may call this region the nuclear cup. Forming the rim of 
the mouth and extending to the conical tip is the capsule, which 
may be regarded as a sort of a conical enclosure around the 
sperm body. 

From the foregoing observations on the series of changes, 
featured by diminution of the constituent materials of the sper- 
matid, each time followed by reorganization and differentiation, 

310 The Philippine Journal of Science *^^ 

it can be said that the spermatozoon is composed of nothing 
more than concentrated nuclear materials and a certain amount 
of cytoplasm packed in a conical capsule. 

Acrohlast and acrosome. — ^The most conspicuous features in 
the later part of spermatogenesis concern the appearance of the 
acroblast and the series of changes that finally lead to its 
transformation into a capsule. Acroblast is a body or group 
of bodies derived from the substance of the idiosome and 
Golgi-bodies and from which arises the acrosome (the capsule 
in this case). A brief summary of the behaviour and move- 
ments of these bodies will serve to portray the role they play in 
sperm formation. 

In these crabs (S. tranquebarica) , the acroblast is at first 
identified as a group of granular bodies (the pro-acrosome gra- 
nules) within the idiosome (Plate 3, fig. 5, p. ac). The idio- 
some is composed of a clear substance called by some authors 
as the "sphere substance.^' From its position at the side, the 
acroblast shifts anterior to that of the nucleus and moves to- 
wards the apical pole (Plate 3, figs. 5, 6), where it appears 
as a heavily stained body inside the idiosome. This implies 
that during interkiiiesis, the granular bodies have disaggregated 
and later some portions reaggregated to form the acroblast 
(Plate 3, fig. 6, ac), while the r^t is thrown off as the (Jolgi- 
bodies remnant (Plate 3, fig. 6, g) . 

In Plate 3, fig. 7, the idiosome bulges out to a conical form; 
there is no indication of the presence of the acroblast within its 
area, except for the presence of some fine granules. Obviously 
the acroblast has undergone disaggregation- 

In Plate 3, fig. 9, the acroblast is shown as a group of heavily 
stained granular bodies near the apical pole. This indicates 
that, during the intervening phases, the acroblast underwent 
disaggregation and now it is again undergoing reaggregation. 

Plate 3, fig. 10, shows the acroblast as two distinct bodies 
opposite each other at the periphery of the idiosome. The sub- 
sequent contraction that takes place brings these two bodies 
together (Plate 3, fig. 11, ac). With the sloughing off of some 
^of the cytoplasm, the more homogenous body which is leaning 
more towards the outside of the idiosome becomes detached and 
sinks towards the base (Plate 3, fig. 12), from where it will 
be cast off eventually. We shall call this the acroblast-rem- 
nant (ac. r). The other acroblast body is not seen in this 
figure; apparently it has disaggregated. 

^^» 3 Estampador: Studies on Spermatogenesis and Oogenesis 311 

In Plate 3, fig, 15, the acroblast reappears as a heavily 
stained body at the border of the idiosome. In the sub- 
sequent phase, it splits into two bodies, one remaining at the 
apical pole to become the definitive acrosome body; the other 
sinking towards the base, perhaps to become Ihe second acrS-* 
blast-remnant and to be eventually cast off. It is more likely, 
however, that its materials may become finely dispersed and 
used in the formation of the nuclear cup (Plate 3, figs* 16, 17) • 

During the series of concentration, contraction, and sloughing 
off of materials, the definitive acrosome body migrates nearer 
to the base (Plate 3, figs. 16, 17) ; then ultimately disaggre- 
gates to become incorporated into the materials that form the 
capsule of the spermatozoon. 

The Capsule. — From the foregoing observations, we can re- 
gard the capsule as the structure formed from the idiosome 
and its sphere substance, together with what is left of the 
acroblast. Hence, it can be considered homologous with the 
so-called acrosome in some sperms, since both structures are 
derived from the same materials. 

Cursory examination of Plate 3, figs. 16, 17, and 18 seems to 
indicate that the direction of the movement of the definitive 
acrosome body is from the base towards the apical pole (Plate 
3, fig. 17). Wilson (1928, p. 381)* states: "the most con- 
spicuous feature in the history of the acroblast is its ultimate 
separation into two parts — one of which moves to the anterior 
pole and there gives rise to the definitive acrosome, while the 
other and often larger portion constitutes acroblast-remnant 
(idiosome-remnant) containing the original Golgi-bodies sur- 
rounding the sphere.'' This seems to be the case only in the 
early stages. 

The observation described by Wilson may really be true 
for some animals. In these crabs, in the later stages, there seems 
to be no such movement, because the definitive acrosome is 
formed in the apical region, the idiosome which holds the acro- 
blast being located in this area. Obviously, the series of re- 
organization and differentiation involving the acroblast and 
evidently the idiosome itself must take place in this zone. Hence, 
in this particular case, the more apparent direction of the move- 
ment of the definitive acrosome is from the apical pole and 
towards the base; whence, it presumably furnishes the material 
that forms the part of the capsule enclosing the nuclear cup. 

312 The Philippine Journal of Science ^^^ 

Capsule formation in other crabs. — Binford (1913)2 made 
the following observations: 

1. The capsules arise in the cytoplasm as a clear vacuole which 
may be stained with Lichtgrun. Its contents is gradually changed 
and have a greater affinity for chromatin stains. 

2. From a granule on the proximal side of the capsule, the central 
body develops into the capsule 

3. The mitochondrial substance is aggregated from the cytoplasm 
and deposited as a ring between the nucleus and the capsule. 

Wilson (1886), Sabater (1893) and Koltzoff (1906) described 
a similar vesicle which arises in the cytoplasm either against 
the nucleus or close to it. The vesicle described by Binford 
and other authors might be the idiosome and the granule re- 
ferred to, the acroblast. 

The observations of the foregoing authors, though made on 
different crustacean decapods, seem to be in agreement with my 
assumption that the idiosome and the acroblast form the cap- 
sule; it cannot be merely from a vesicle, this term being merely 
descriptive, indicating a structural feature. The idiosome, on 
the other hand, has in its make-up something that has active 
directive influence. Perhaps, it is this unknown factor that 
influences the transformation of the capsule during the pene- 
tration of the sperm into the egg. Binford himself (1913) « 
said: **for the interpretation of the entrance of the sperma- 
tozoon into the egg, the transformation of the capsule is more 
important than the changes in the protoplasmic cup.'' 

II. Spermatogenesis in the class without tetrad formation 
(S. tranqvsharica) . — ^The mode of spermiogenesis involving cells 
with no tetrad formation differs markedly in many aspects 
from that of the class in which there is- In the former, the 
center of activity seems to be localized in the nucleus and to a 
very limited area immediately around it. The remaining por- 
tion of the cell apparently remains passive. The phases of 
changes are not so picturesque, and there is an obvious attempt 
at shortening the period of sperm formation. A brief descrip- 
tion of some of the features may help show the differences 
between these two types of spermiogenesis. 

Plate 4, fig. 1, shows a growing auxocyte, drawn from the 
cross-s^tion of a follicular tubule. As in the auxocytes with 
tetrads, the characteristic features are: (1) the big nucleus 
which is basal in position; (2) the greater amount of cytoplasm 
present, compared with that in the nurse cells; (3) hetero- 
pycnosis, that is, the condensation of some chromatin materials 

^^' ^ Estampador: Studies on Spermatogenesis and Oogenesis 313 

to form the chromosome-nucleolus. The nucleus, however, is 
relatively smaller in size compared with that of an auxocyte 
belonging to the class with tetrads. 

Plate 4, fig 2, is drawn from the surface view of an auxocyte 
in a later stage. The chromatin materials are now organized 
into chromatids. There is no indication of tetrad formation. 

Plate 4, fig 3, is drawn from the longitudinal section of the 
follicle. The condition of the nucleus of the auxocyte indicates 
that some sort of rearrangement is taking place; perhaps this 
foreshadows the differentiation of the spermatid body. 

In the class with tetrad formation, we differentiate a pro- 
spermatid stage which subsequently undergoes a series of trans- 
formations involving the elimination of a considerably large 
amount of cytoplasmic materials prior to the formation of the 
definitive spermatid body. On the other hand, in the class with 
no tetrads this series of changes is omitted; hence, the mode 
of spermiogenesis differs in that it is somewhat abbreviated. 

The chromosomes (Plate 4, fig. 4) ) are very much smaller 
compared with those present in the auxocytes with tetrad 
formation (Plate 3, figs. 4, 5, and 7). In both classes, how- 
ever, the chromosomes are segmented. The hetero-chromosome 
may be seen as V-shaped body (h). Besides the relatively 
smaller size of the chromosomes, another significant difference 
is in number— -there are only five, instead of six, (Plate 3, fig. 
13 and Plate 4, fig. 9). This is a case of digamety. The dif- 
ference in number is due to the presence in the group having 
tetrads, of an extra chromosome, the monosome (Plate 3, figs. 

In Plate 4, fig. 5 are shown two spermatids that have differen- 
tiated from their respective auxocytes. In (a) the acroblast 
(ac) appears as a heavily stained body at the apical pole and 
the rather large nucleus is at the base. In (6) the acroblast 
has evidently split into two bodies; the one inside the acroblast- 
vesicle {v. ac} within the idiosome will probably form the 
definitive acrosome body; the other is not shown. No pro-acro- 
some granules have been observed in this case (Plate 3, fig. 5) ; 
the acroblast suddenly appeared as a single chromophyllic body 
within the acroblast-vesicle. The reticular condition of the 
auxocyte body may be attributed to the disorganization of the 

The darkly stained masses (no) are evidently the sloughed- 
off chromatin materials and the other similarly stained mass (g) 
presumably represents the Golgi-remnant bodies. These masses 

314 The Philippine Journal of Science ^^^ 

have their counterpart in Plate 3, fig, 6, It can be inferred that, 
with the disappearance of the nuclear wall as the chromosomes 
are formed, some of the chromatin materials are discarded. In 
other words, chromatin diminution took place when the sper- 
matid differentiates from the auxocyte body; that is, during 
the differentiation, and in the reorganization that followed, 
only so much chromatin material is retained as is proportionate 
to the cytoplasm of the spermatid body. In other words, the 
amount of cytoplasm incorporated into the spermatid body 
determines the quantity of chromatin material that is to b^ 
included. This premise, however, does not necessarily preclude 
the occurrence of a reverse condition. This brings up the 
question regarding the casual relation between the nuclear 
volume and cytoplasmic growth — ^the so-called "karyoplasmic- 
ratio" of R. Hertwig (1903, 1908, etc,)- 

The present case seems to be in line with Conklin*s observa- 
tion that **the nuclear volume varies not with the total volume 
of the cytosome but with that of its active protoplasm," Com- 
parative measurements show a striking correspondence in size 
between the definitive spermatid body and the nucleus of the 
auxocjrte at the termination of the growth period. The resem- 
blance is such that on cursory examination the spermatid is 
likely to be mistaken as the nucleus. 

This mode of spermiogenesis (in auxocytes without tetrads) 
was previously described as characterized by the apparent local- 
ization of activities in the nucleus including certain limited 
portion of the cytosome immediately around it. During the 
differentiation of the spermatid from the auxocyte body, the 
nucleus underwent reorganization so as to make its mass rel- 
atively proportional to that of the active protoplasm which is 
to be incorporated into the spermatid body. This perhaps ac- 
counted for the sloughing off of some chromatin materials. 

There are features that yet remain to be explained, such as: 

(1) In the class that has no tetrad formation, what limit the 
activities to a very restricted portion of the auxocyte body? 

(2) Why does the whole auxocyte body take active part in the 
transformation in the other class (with tetrads)? 

Aside from the physical constitution of the chromosomes, the 
particular phase wherein the two classes differ conspicuously 
is in the manner by which their respective spermatids dif- 
ferentiate from the auxocjrte. Otherwise, in some broad aspects, 
the other phases of transformation are discernibly parallel. 

78, ^ Estampador: Studies on Spermatogenesis and Oogenesis 315 

Plate 4, fig. 6, presumably represents a later phase following 
the contraction of the spermatid body represented in Plate 4, 
fig. 5 (&)• The spermatid body is now elongated and the de- 
finitive acrosome occupies a central position within the idiosome 
at the apical pole, while the acroblast-remnant (ac. r) lies at 
the periphery of the base, eventually to be cast off. 

In one of the subsequent phases, a figure resembling a lamp- 
shade is formed (Plate 4, fig. 6), with the acrosome body at the 
apical area. (Cf. Plate 3, figs. 10 and 11, which belong to the 
group with tetrad formation.) 

The next phase is featured by the extension of the apical 
area to a narrow tip while the base describes a flange at the 
periphery, (Plate 4, fig. 8). 

Plate 4, fig. 9, shows the basal portion of a spermatid. There 
are five chromosomes. (Cf. Plate 3, fig. 13.) 

Plate 4, fig. 10, shows a spermatid in about the last phase of 

The existence of digamety in the sperms of this group of crabs 
iS. tranqmbarica) is unquestionably evident. Whether dime- 
galy also exists at the same time is yet to be proved. 


There is only one method of spermiogenesis in this group, 
in sharp contrast to that in S. tranquebarica. Some of the 
phases of transformation are shown in the following illustra- 

Apparently, after the growth period the auxocyte nucleus 
splits into two spermatid nuclei (Plate 5, fig. 3). During the 
subsequent reorganization, some of the cytoplasmic materials 
around each daughter nucleus become incorporated into the 
spermatid body (Plate 5, fig. 3). The portion of auxocyte 
which is involved in the process of spermatid differentiation 
of the cytosome includes the zones in the immediate vicinity 
of the spermatid nuclei and the area around the rather enor- 
mously large centrosome (a), wherein a centriole can be seen 
at the center. 

Plate 5, fig. 5, represents a pair of spermatid bodies that have 
developed from the spermatid nuclei referred to above. Ap- 
parently they are in the contraction phase; the dark condensed 
chromatin mass, evident in each of them, represents the sex- 
chromosomes. Each spermatid eventually transforms into a 
spermatozoon through a series of differentiation and reorganiza- 
tion accompanied by the diminution of both the cytoplasmic and 
karyoplasmic materials. 

316 The Philippine Journal of Science ^^^^ 

That the two daughter spermatid nuclei transform into 
functional spermatozoa is quite evident. Spermatids in about 
the same phases of transformation are commonly seen very 
close together (Plate 5, fig. 5) and spermatozoa in pairs are 
not uncommon. In this figure the two spermatids had evidently 
undergone diminution of both cytoplasmic and karyoplasmic 
materials. Both are evidently in the sinizesis stage. 

Plate 5, fig. 6, shows a side view of a spermatid in a later 
phase. The acroblast appears as a big black mass at the peri- 
phery of a rather large idiosome (t). 

The subsequent contraction splits the acroblast into two por- 
tions, one sinking to the base as the acroblast-remnant (Plate 
5, fig. 7, ac. r), and the other remaining at the apical area as 
the definitive acrosome body. The figure shows the spermatid 
from the top, which is narrow, while the base is expanded. The 
figure corresponds to what we previously termed as the lamp- 
shade phase, (in S. tranquebarica, Plate 3, fig. 10 ; Plate 4, fig. 7) . 

Plate 5, fig. 8, represents an example of spermatozoa in pairs. 
Their presence in close proximity to each other is taken as 
an indication that the two spermatid nuclei that differentiate 
from the nucleus of an auxocyte may transform into two 
functional spermatozoa. In this figure, one of the pair is seen 
from the basal-side view; only the basal portion of the other 
is shown. Both spermatozoa show the central position of the 
nucleus in the nuclear cup. 

Histogenesis in this group differs in many aspects from both 
of the two modes of spermiogenesis that we observe in S. tran-- 
quebarica. In the initial phases: (1) no tetrad formation is 
seen; (2) no pairing of chromosomes; (S) only a limited portion 
of the cytosome is actively involved. These are in sharp contrast 
to the mode of spermiogenesis involving the class of auxocytes 
(with tetrad formation) in S. tranquebarica, wherein the whole 
cell participates actively. 

Compared with the mode of spermiogenesis that takes place 
in the class without tetrad formation, the contrast is also 
obvious. In S. tranquebarica, the nucleus of an auxocyte dif- 
ferentiates directly into a spermatid; presumably, this takes 
place by incorporating some of the cytoplasm after the sloughing 
off of a certain amount of the karyoplasm. In S. oceanica, on 
the other hand, the auxocyte nucleus splits first into two 
spermatid nuclei; then each nucleus reorganizes into a sper- 
matid by incorporating a certain amount of the surrounding 

^8» 3 Estampador: Studies on Spermatogenesis and Oogenesis 317 

However, in the later phases of spermatid-transformation, 
the above mentioned three quite distinct modes of spermiogenesis 
run parallel in several aspects as indicated by: (1) the series 
of concentration, contraction and sloughing off of some materials 
with the consequent diminution of the size of the spermatid, each 
time followed by differentiation and reorganization until the de- 
finitive spermatozoon form is attained; (2) the behavior and 
movements of the acroblast and acrosome are essentially similar. 

The chromosomes (Plate 5, fig. 9) of S. oceanica, exhibit the 
tendency of being formed loosely by groups of twos, threes 
(triosome or triplicate), and fours (quadruplicate) sets of chro- 
matids. This is in contrast with the distinct-segmented char- 
acter of the chromosomes of S, tranqueharica. (Plate 3, figs. 
4, 5 and 7.) 

The haploid number of chromosomes in S. oceanica is six 
(Plate 5, fig. 10). There is only one mode of spermiogenesis, 
in contrast to the two methods of sperm formation in S. tranque" 


Histogenesis in this group closely parallels that of S. tranque- 
barica in the sense that there are two types: (1) in one mode of 
spermiogenesis, the whole cell is practically involved actively. 
This is indicated in Plate 5, fig. 11, which shows a pro-spermatid 
in the contraction phase with some of its csrtoplasmic and 
chromatin materials being sloughed off. The size is relatively 
large compared with that of the kinoplasmic area indicated in 
Plate 6, fig. 1. 

We need not follow closely the details of the changes. On 
the whole, it can be said that, the phases of transformation 
broadly resemble in many essential details the spermiogenesis 
in one class of auxocytes (with tetrad formation) in S. tran- 

(2) The other type involves some broad aspects which suggest 
a mode of spermatid-diff erentiation somewhat intermediate be- 
tween that in S. oceanica, on one side, and that in S. tranque* 
barica (only in auxoc3rtes without tetrads), on the other, that 
is, if we are to base the condition on the kinoplasmic area. 
There seems to be a degree of gradation regarding the relative 
amount of the cjrtoplasm which is actively involved — ^practically 
nil in S. tranquebarica (only in auxocytes without tetrads), 
small in paramamosain, and considerably greater in S. oceanica. 
(Cf. Plate 4, fig. 5; Plate 6, fig. 1 and Plate 5, fig. 3.) 

318 The Philippine Journal of Science ^^^ 

In paramamosain, the nucleus with some portion of the 
eytosome (Plate 6, fig. 1) differentiates from the auxocyte and 
splits into two, each half transforming into a spermatid. The 
mode of spermatid-differentiation in this group is different then 
from what were observed in the two other groups (S. tranque- 
barica and S. oceanica). 

Frequently, two spermatozoa can be seen side by side within 
the remains of an auxocyte. This indicates that the two 
spermatozoa must have developed from two spermatids, like 
the two referred to above. Plate 6, fig. 2, represents one of 
such pairs; the two spermatids have already attained the de- 
finitive form of the sperm; apparently, they have to undergo 
reorganization before they become full-fledged spermatozoa. 
The reticular appearance of the auxocyte body is noteworthy; 
the same condition may be olBerved in Plate 4, fig. 5. Perhaps, 
this condition is due to the withdrawal of the eytosome from 
some parts of the cell during the time when the spermatids are 
differentiating from the auxocyte body. Presumably, the large 
clear space (Plate 6, fig. 2, c) above the spermatids represents 
the centrosome with some centrioles and centroplasm. 

It is not unlikely to find two types of spermatozoa in the 
variety paramamosain, since there are two modes of spermio- 
genesis. This condition is observed in S. trwnqueharica. 


Spermiogenesis in this group proceeds in two distinct fashions 
varying distinctly in some details from the types that are 
observed in the preceding groups. In this species, the sper- 
matids may differentiate from the auxocytes by either of two 
quite distinctly different methods. 

I. In one class, after the growth period, the auxocyte nucleus 
undergoes a somewhat general reorganization. This involves 
both karyoplasmic and chromatin materials and their subsequent 
diminution (Plate 6, figs. 7, 8 and 9). The spermatid so 
differentiated then underg(^s transformation. 

Plate 6, fig. 7, shows an auxocyte nucleus apparently in the 
phase of the series of changes that lead to its transformation 
into a spermatid body. The chromosome-nucleolus appears as 
a condensed mass, while the other chromosomes are visibly 
still somewhat in a reticulated condition. The karyoplasmic 
materials are more or less concentrated on the opposite pole. 
The centrosome (c) appears as a clear space containing some 
centroplasm. With acid fuchsin, anilin blue and haematoxylin, 
the latter appears as a somewhat dark mass with a bluish tint. 

^*» 3 Estampador: Sttidies on Spermatogenesis and Oogenesis 319 

In the subsequent contraction phase, some of the materials 
are sloughed off. Plate 6, fig. 8, shows a picture wherein the 
process is apparently going on. The karyoplasm is in the 
diffused condition and the chromatids, excepting those that 
constitute the sex-chromosomes, are aligned along the peri- 
phery of the differentiating body. 

Plate 6, fig. 9, shows a phase of reorganization following the 
differentiation and the subsequent sloughing off of some ma- 
terials. The spermatid nucleus is in the netlike stage or 
resting period and the chromosomes are temporarily in a 
reticulated condition. The sloughed-off materials (rp) are seen 
above it. 

Spermatids differentiated in this manner seem to be less 
common than the other class and, in their later phases of 
transformation, have not been followed very closely. This is 
quite difficult, because the determination as to how a spermatid 
differentiates from the auxocyte is rather uncertain. It is quite 
true that the spematids belonging to the other class usually 
occur in pairs and, hence, they can be reorganized easily; but 
there still remains the uncertainty due to the accidental separa- 
tion of the pairs which may happen during the sectioning of the 

II. The other form of spermiogenesis in this species is char- 
acterized by the differentiation of the auxocyte nucleus as a 
body to form the spermatid nucleus (Plate 6, fig. 10). This 
then proceeds to reorganize and ultimately splits into two. 
Perhaps this explains the fact that many spermatids and sper- 
matozoa are in pairs. Plate 7, fig. 4, shows a pair of spermatids 
still within the auxocyte body. The photomicrographic picture 
of a spermatocyst (Plate 6, fig. 6) also illustrates this condition. 

The process of division is featured by what we may term 
"progressive heteropycnocis." The chromosome-nucleolus ap- 
pears to be the focal point around which the activities are more 
obvious. The following figures illustrate some of the more 
characteristic features. 

Plate 6, fig. 10, shows the nucleus of the auxocyte at the 
termination of the growth period. Apparently the resting 
phase, which is indicated by the netlike stage, or resting 
condition, in which all the other chromatin materials are retic- 
ulated; they are being broken by the initiation of activities. 
This can be inferred from the apparent tendency of the nu- 
clear material to concentrate around the chromosome-nucleolus. 

320 The Philippine Journal of Science *^* 

Obviously this results in the realignment of the nuclear con- 
stituents which finally segregate into two portions (Plate 6, 
figs. 11, 12, and Plate 7, fig- 1). 

Plate 7, fig. 2, shows the nucleus only, wherein the chromo- 
some-nucleolus (presumably the XX-chromosomes) now elon- 
gates at the equatorial region. The doubled appearance and the 
presence of clefts at both ends, instead of the two ends taper- 
ing and pointed, as observed by some authors in other animals, 
indicate that, in this particular case, the chromosome-nucleolus 
is composed of the XX-chromosomes. Obviously the pair is in 
diakinetic position, preparatory to their separation; hence, each 
daugther nucleus will have one X-chromosome. 

The process of division is shown in Plate 7, fig. 3. A peculiar 
feature is seen in the sudden appearance of presumably the 
idiosome (t) at the angle where the splitting takes place. 

Basing on what has been observed regarding the role and 
the active participation of the idiosome in the process of trans- 
formation in the other groups, it can be inferred that in this 
case a portion of the idiosome is incorporated into the body 
of each spermatid during the reorganization that follows the 

This view is apparently supported by the appearance of the 
acroblast in a later phase (Plate 7, fig. 5). The acroblast 
appears at the apex of a cuplike structure; the nucleus fills 
the mouth of the cup. In this connection, it may be pointed 
out that the features and the circumstances associated with 
the apperance of the idiosome and the acroblast are different 
from what have been observed in S. tranqmbarica. 

The features are also different in the two other groups 
(S. oceaniea and in the variety paramamosain) , where the aero* 
blast appears suddenly within the idiosome in the apical pole 
of the spermatid; whereas in this species (S. serrata)^ the 
idiosome appears first in the cjrtoplasm of the auxocyte at the 
place where the nucleus is splitting. On the whole, however, 
in the last three groups, namely: S. oceaniea, the variety 
paramamosain, and S. serrata, the appearance of the acroblast 
takes place at some later stage of the transformation. 

Plate 7, figs. 6 and 7, represent spermatids in the later stages. 
Plate 7, fig. 8, shows a spermatozoon. The existence of two 
classes of spermatozoa in this species is a possibility that can 
be inferred from the two distinctive ways by which spermatids 
differentiate from the auxocyte. 

^^ 3 Estampador: Studies on Spermatogenesis and Oogenesis 321 


The principal points brought out in this study of sperma- 
togenesis in these four groups of crabs are the following: 

!• The lumen of the follicular tubule is lined by a single 
layer of cells. These cells are of three kinds: (a) the spermato- 
blasts; (6) the nurse cells; (c) the secretory cells. 

2- The spermatoblasts (auxocytes) grow at the expense of 
the nurse cells and eventually develop into spermatozoa. The 
cells of the gonads of these four groups show discernible dif- 
f erences in size, form, and in their structural make-up. Gen- 
erally, the spermatoblasts develop singly, but in some cases 
several cells may form into a group enclosed by a cyst. This 
group formation is called spermatocyst. Spermatocyst forma- 
tion occurs quite commonly in S. serrata and its variety para- 
mamosain, but quite rare in S* tranquebarica. It does not 
seem to occur in S. oceanica (banhawin). 

3. The ways by which spermatids differentiate from the 
auxocyte differ in some broad aspects: 

a. In S. tranqueharica, it proceeds in two ways: 

(1) First, there is a pro-spermatid stage, which is char- 

acterized by the sloughing off of considerable amount 
of both cytoplasmic and karyoplasmic materials, lead- 
ing to the formation of the definitive spermatid body. 
There is tetrad formation. The haploid number of 
chromosomes is six. 

(2) Second, the nucleus, perhaps with a certain amount of 

cytoplasm immediately around it, differentiates into a 
spermatid. There is no tetrad formation and the 
haploid number of chromosomes is five. Therefore, 
the sperms of S, tranquebarica exhibit digamety. 

b. In S, oeeaniea, spermatid formation proceeds in only one way. 

The auxocyte nucleus first splits into two spermatid nuclei; 
then each daughter nucleus reorganizes by incorporating 
some of the cytoplasm to form into a definitive spermatid 
«, In the variety paramamosain, spermatid formation proceeds in 
a somewhat parallel manner to that in S. tranquebarica, in 
the sense that there are also two methods, namely: 

(1) In one type, as in S. tranquebarica, there is a pro- 

spermatid stage; that is, the formation of the defin- 
itive body is preceded by gradual diminution of both 
the cytoplasm and nuclear materials. 

(2) In the other type, the auxocyte nucleus differentiate! 

as a body from the auxocyte by forming an active 
kinoplasmic area in which only a very limited amount 

322 ^^^ Philippine Journal of Science *^* 

of the cytoplasm is incorporated. This active mass 
soon splits into two, forming two spermatids, 
d. In S. serrata, spermatids may differentiate from the auxocytes 
by two methods: one resembles somewhat the mode of 
spermatid differentiation (only auxocytes without tetrad 
formation) in S, tranquebarica, but rather more abbre- 
viated; this is rather obscure and needs to be studied further. 
The other method parallels that of S. oceanica^ but in a 
fashion that is somewhat abridged and involves only con- 
siderably less cytoplasm. 

(1) In the first case, the auxocyte nucleus is transformed 

into a spermatid through a process of differentiation 
and reorganization during which some of the ma- 
terials are sloughed off. Apparently, if at all, only 
one spermatozoon develops from the auxocyte. 

(2) In the second, the nuclear materials undergo a sort 

of realignment distinctly different in features from 
those observed in the first. The nucleus then divides 
into two spermatid nuclei, and each daughter nucleus 
reorganizes to form into a spermatid. Evidently, 
two spermatozoa are produced. Progressive hetero- 
pycnosis is a very conspicuous feature in this type of 
spermatid formation. 

4. In all groups the main features of the transformation 
of spermatids to spermatozoa involve a series of processes of 
concentration, contraction, and diminution of both of the cyto- 
plasmic and nuclear materials including other cellular bodies. 

5. One main interesting feature involves the participation 
of the idiosome, the appearance and movement of the acro- 
blast body tending toward the subsequent formation of the 

6. The general form of the sperms is conical; the broad 
basal portion that holds the nucleus is the nuclear cup, the 
rim of the cup and the conical portion constitute the capsule. 
This latter is apparently composed of the idiosome which 
holds the non-chromophyllic "sphere-substance", and the acro- 
blast material. The differences in size and in general outline 
between the spermatozoa of these four groups of crabs, and 
even between the spermatozoa of the same variety, are quite 
discernible. These differences, however, are hard to determine 
accurately due to several causes, such as: (1) the difficulty of 
getting accurate observation because of their being very minute ; 
(2) the variable factors involved in staining and destaining, etc. 


The abdomen is broad and all the segments are freely mova- 
ble, (Plate 1, fig. 4). There is a pair of appendages on each of 

'8' 3 Estampador: Studies on Spermatogenesis and Oogenesis 323 

the second, third, fourth and fifth somites. The external genital 
apertures (Plate I. fig. 1, g. o) are a pair of large openings 
situated on the sternum of the sixth thoracic somite. The sterna 
are not so concave as those of the male, and the abdomen is not 
so closely applied to the thorax. 

The ovaries (Plate 1, fig. 2) are paired, and lie in a position 
similar to that of the male reproductive organs. There is, 
however, a considerable posterior prolongation of each ovary. 
As in the male, the two anterior lateral portions are connected 
behind the pyloric f oregut, the connection forming a sort of 
a bridge over the midgut. Behind this transverse connection, 
each ovary is prolonged backwards as a narrow strip extending 
to the extr^ne posterior end of the thorax, where they fuse 
together in mature specimens. These posterior extensions of 
the ovaries occupy a similar position to that of the vasa de- 
ferentia of the male; that is, they lie above the hindgut and 
below the pericardium. Beneath the anterior portion of the 
latter, each posterior branch is connected on its outer side with 
a large sac, the spermatheca (spt). This continues as a short 
oviduct which opens to the exterior by means of the vulva on 
the sternum of the sixth thoracic somite. 

The seminal receptacle is composed of two parts, namely: 

(1) the glandular portion which is contiguous with the ovary, 

(2) the portion lined with chitin which is continuous with the 
oviduct. The oviduct (Plate 1, fig. 2, od) is chitinous, its ter- 
minal portion expanded to form a sort of funnel-like structure 
leading to the vulva. 

The condition of the ovary depends upon the degree of 
maturity of the ova. When immature, the gonads are small 
and pale, and the presence of the eggs cannot be detected by 
the naked eye. But when mature, the gonads fill up almost 
the whole dorsal surface of the cephalothorax. They are orange- 
red and the eggs can be readily distinguished. The red color 
is due to the presence of a large amount of yolk. 

The ovary is composed of numerous follicular tubules. When 
immature, the general outline in cross-section (Plate 1, fig. 3) 
is that of a triangle, with the apex fitting into the stemo-cara- 
pace angle. With the ripening of the eggs, the ovarian walls 
become very much distended; the outline of the gonads at this 
time becomes irregular. The gonads fill up the stemo-carapace 
angle and practically covering up the entire dorsal surface of 
the branchial r^on beneath the hypodermis. The follicles 
are no longer distinct; under the microscope, only some portions 


324 The Philippine Journal of Science *^* 

of the follicular walls can be seen. The epithelial cells are 
obliterated; the gonads being filled up with compact masses of 

The cavity of the follicular tubules (Plate 7, fig. 9) is lined 
by a single layer of epithelial cells. These ceUs may be grouped 
under four categories, namely: (1) ooblast; (2) nurse cells; (3) 
the secretory cells and (4) the auxiliary secretory cells. 

The secretory cells. — Compared with those present in the 
testicular follicles, the ovarian secretory cells are generally 
bigger and more numerous. In a young ovarian follicle, there 
seems to be no marked differences between the principal and 
the auxiliary secretory cells, as both groups are vacuolated, 
the only difference being in the presence of a large vesicle in 
the former. This vesicle begins to enlarge (Plate 7, fig. 9, c. c), 
the enlargement seemingly correlated with the dissolution of 
the materials of the adjacent cells. In other words, the ad- 
joining vacuolated cells act as auxiliary secreting cells by yielding 
their materials for the production of the metaplasm of the 
principal secretory cells. Presumably, there must be an enzjnme 
the action of which is responsible for the conversion of cellular 
materials into secretory products. This metaplasmic product 
takes up a bluish color with iron-haematoxylin, acid-f uchsin and 
anilin stain combination. 


The ooblast (auxocytes) develop into eggs; that is, the epi- 
thelial cells metamorphose directly into ovocytes without divi- 
sion. The auxocytes can be easily recognized from the nurse 
cells by their relatively bigger germinal vesicles, and the pre- 
sence of more cytoplasmic materials. In a young auxocyte, 
the cytoplasm is at first small in amount and simple in structure, 
seemingly consisting almost wholly of optically homogeneous 
hyaloplasm. As growth proceeds, Ihe appearance becomes more 
or less alveolar, and there is an enormous increase in the am- 
ount of ooplasmic substance. In well-stained sections, stream- 
ers of both karyoplasmic materials and cytosome from the 
neighboring cells are discernible. The growing auxocyte does 
not only receive material contributions from the neighboring 
cells, but in some cases it engulfs a whole cell. Vestiges of 
such engulfed cells are quite apparent in some full-grown au- 

The description above is based on the auxocytes of S. oceanica 
and those of 5. tranqueharica. The auxocytes of the variety 

'*' 3 Estampador: Stvdies on Spermatogenesis and Oogenesis 325 

paramamosain and especially those of S. serrata appear to be 
smaller and less alveolar; hence, they appear more compact. 

The process of oSgenesis parallels that of spermatogenesis 
to a certain extent. Both processes are featured by the growth 
of the auxocytes at the expense of the nurse cells; likewise, 
both germ cells undergo transformation at the termination of 
their growth period. 

Odkinesis. — ^The amitotic ookinesis through which the ovo- 
cytes pass is apparently less complicated, compared with the 
changes that feature spermiogenesis. And, like the latter, the 
phases of transformation in each of the four groups differ 
in many aspects, some features being distinctive within the 


This proceeds in somewhat similar fashion as spermiogenesis 
with respect to the extent of the kinoplasmic area involved, 
namely: (1) in one type, the whole cell is actively involved; 
(2) in the other, only the nucleus and the area immediately 
around it participate actively. These two modes of ookinesis 
produce ovocytes that show differences in size, and also in the 
constitution of their chromosomes. The latter aspect, however, 
seems to be temporary in nature. These two forms of ovocyte 
are referred to as the macro and the micro-type. 

The following figures illustrate some of the characteristics 
that feature the development of the macro-ovocytes. 

Plate 7, flg. 10, shows the surface view of two developing 
auxocytes and Plate 7, fig. 11, represents one from the cross- 
section of a follicular tubule. The disorganized state of both 
the cytosome and the nuclei of the nurse cells is an indication 
that some of their materials are being absorbed by the growing 

Plate 8, fig. 1, represents an ovocyte at the termination of the 
growth period. The germinal vesicle has grown to almost 
four times its original size. It seems to be in the zygotene stage. 
The pairing of the spireme is quite obvious. Around the ger- 
minal vesicle, roughly in the form of a horse-shoe, is the pallial 
layer, or vitellogenous mass (v; also Plate 12, fig. 4). The 
Golgi-bodies are seen as the heavily stained masses scattered in 
this layer. Vestiges of a nurse cell are discernible. 

In Plate 8, fig. 2, the size of the ovocyte and its germinal 
vesicle has decreased considerably. Perhaps this is because 
of the contraction and the subsequent sloughing off of some 
of the cellular materials. The condition of the germinal vesicle 
is apparently that of the pachytene stage, some pairs of spireme 

326 The Philippine Journal of Science ^^ 

having fus^ and b^ame thicker threads. The tetrad forma- 
tion is evident in both Plate 8, figs* 1 and 2, as shown by the 
alignment of chromatids into groups of four. The vitellogenous 
layer is no longer present, but instead, irregular spherule masses 
that stain quite deeply at the periphery are scattered in the 
cytoplasm. Presumably, these masses foreshadow the formation 
of yolk-masses that develop later. Apparently these masses 
became finely disseminated through the ooplasm, since they do 
not appear in the stag^ immediately following. 

Plate 8, fig. 3, shows the chromosomes scattered in the homo- 
geneously diffused karyoplasm. There are apparently twelve, 
excluding the supernumerary ones. The mai^^ containing 
deeply stained bodies are presumably composed of extruded 
nuclear materials and some Golgi-bodies. The XX-pair (h) is 
conspicuous. Apparently the germinal vesicle is undergoing 
an ungual division, leading to the formation of the first polar 
body; and the extrusion of some nuclear material takes place 

Plate 8, fig. 4, shows an ovocyte with the polar nucleus being 
extruded out, together with some cytoplasmic material. It is 
in the contraction phase. 

It seems that the ovocyte after the extrusion of the first polar 
body either goes through a temporary resting stage (Plate 8, fig. 
5) or proceeds in its development. The latter case is indicated 
in Plate 8, fig. 6. The formation of chromatids in groups of 
twos indicates the dyad formation and their alignment with re- 
spect to the other pairs implies that they are in parasynaptic 
association. The six chromatids in a circle compose the hetero- 
chromosomas (fe), the XX-pair in this case, since they are more 
or less alike. The monosome (m) is shown as a heavily stained 
body close to the nuclear wall. 

Plate 8, fig. 7, shows the chromosomes prior to the second 
meiosis. The hetero-chromosomes (h) are easily recognizable, 
as is the monosome (m) which has no pair. Presumably the 
clear area in the vicinity of the germinal vesicle is the attraction 
sphere (a. s) and the two slightly stained masses within are 
the division centers (centrioles). 

It can be inferred from Plate 8, fi^. 7 and 8, that the forma- 
tion of the second polar body apparently proceeds in this man- 
ner: (1) the division centers move apart, followed by the cons- 
triction of the attraction sphere; (2) the nuclear materials 
then become disseminated around the two division centers and 

^«» 3 Estampador: Studies on Spermatogenesis and Oogenesis 327 

(3) around each division center a nucleus is reformed. One 
will become the second polar body. 

In Plate 8, fig. 9, the two daughter nuclei have separated; 
evidently the smaller is the second polar body (p. 62). In the 
subsequent contraction, these two bodies are pulled apart (Plate 
8, fig. 10). The second polar body is distinctly recognizable 
by the presence of several chromatids that are irregularly 
disposed. The egg nucleus is bigger, with the chromosomes 
aligned along the nuclear wall. The expulsion of the polar 
nucleus will probably take place at the termination of the con- 
traction phase. 

The division that has taken place cannot be considered as an 
equational, but reducing division, since qualitatively dissimilar 
halves are produced. The first polar division may also be so 
characterized. Hence, the maturation division in this crab is an 
exception in the sense that ordinarily one maturation division 
is supposed to be equational and the other reductional. 

Ookinesis in the micro-type. — ^The mode of ookinesis that leads 
to the formation of the micro-ovocsrtes is seemingly less involved, 
but the changes are rather obscure. The activity appears to be 
localized in the germinal vesicle, involving only that portion of 
the cytosome immediately around this body. In other words, 
the kinoplasmic area includes only these portions of the auxocyte. 

From Plate 9, figs. 1 and 2, we can infer that at the termina- 
tion of the growth period, the germinal vesicle with some por- 
tion of the cytosome differentiates as a body from the auxocyte 
to form the ovocyte. In this mode of odkinesis, there is seem- 
ingly an attempt at shortening the process of transformation. 

Plate 9, fig. 1, shows an ovocjrte that has differentiated from 
the auxoc3rte, with the first polar body (p. b 1) already formed. 
The deeply stained bodies scattered in the cytosome are probably 

In Plate 9, fig. 2, the polar nucleus is already extruded. 
Evidently the number of chromosomes is six, but they are biva-- 
lent. The heterochromosome (h) is easily recognizable, being 
the biggest and broadly V-shaped. 

The second polar body. — ^The formation of the second polar 
body in the micro-ovocytes takes place after the completion of 
the growth period and after the sperm has penetrated into the 
germinal vesicle of the ovum. This fact is borne out by the 
presence in the ovary of full-grown crabs of germinal vesicles 
which are about twice as big as the germinal vesicles of the other 
groups. Plate 10, figs. 1 and 2, show the relative sizes of the 

328 The Philippine Journal of Science ^^* 

germinal vesicles of the two classes of ova, the macro-ovum and 
the micro-ovum, respectively. The dyad formation is still re- 
cognizable in Plate 10, fig. 2. Another evidence is seen in the 
presence of polar bodies still attached to the eggs that are al- 
ready glued to the endopodite setae (Plate 13, fig. 2). Since 
the eggs are already in advanced stages of development, it is 
quite apparent that the formation of these polocytes must 
have taken place in already mature eggs, or perhaps even later 
when these eggs are already outside the body and undergoing 

Quite frequently some protozoa, especially stalked ciliates, 
attached themselves to the eggs. Their reaction to the fixatives 
is to contract and round up ; hence, they are likely to be confused 
with the polar bodies. They can be distinguished easily, how- 
ever, by their elongated and deeply stained nuclei; while the 
polar bodies have clear nonstaining rounded vesicles, and in 
most cases, the protoplasmic connection between a polocyte 
and an ovum is still visible. 


From the time of the formation of the polar nucleus the egg 
increases immensely in size, from about 29 microns to over 266 
microns in diameter. Undoubtedly this great increase in bulk 
represents a considerably long period of growth. Plate 8, fig. 
10, and Plate 9, fig. 3, give a comparative idea regarding the 
increase in bulk of the ovum. 

The appearance becomes alveolar because of the presence 
of closely crowded yolk-spheres suspended in the hyaloplasm, 
in which also lie numerous smaller granules of various kinds, 
some fatty materials and pigment granules. The ovary then 
becomes more compact and assumes a bright orange-red color 
and there are no longer recognizable traces of either the secre- 
tory or the nurse cells. These cells which originally consti- 
tute the main bulk of the follicle must have been used up to 
furnish materials for the growth of the ova. 

The position of the germinal vesicle is invariably near the 
center, in the young ova and also in older ones which happen 
to be free from pressure. In closely crowded ova, the ger- 
minal vesicles become eccentric in position. The wall of the 
germinal vesicle is generally distinct in most places, except at 
points where it merges with the cytoplasm and from which 
radiating protoplasmic streamers branch outward to the peri- 
pheral cytoplasmic layer (blastema) around the egg. These 

^^*^ Estampador: Studies on Spermatogenesis and Oogenesis S29 

protoplasmic strands course deviously through the spaces be- 
tween the yolk-spheres, and they form the connecting links be- 
tween the germinal vesicle and the rest of the egg. The eggs 
are distinctly centrolecithal. 

The foregoing description is based upon the eggs of S, tran- 


There is only one mode of ookinesis in this species. It pro- 
ceeds in a manner somewhat similar to the mode of ookinesis 
that produces the micro-ovocytes in S. tvanqueharica. There 
are distinct differences, however, as follows: (1) the bulk of 
the auxocyte; the first polar body {p. hi) is already expelled, 
atively much greater in the auxocytes of S. oceanica; (2) hence, 
the ovocytes that develop are relatively bigger (Plate 9, fig. 
2, and Plate 10, fig. 8) ; and (3) the formation of the second 
polar body follows immediately that of the first polocyte. In 
contrast with S. tranqtiebarica which exhibits sexual dimegaly, 
S. oceanica produces only one class of eggs. 

Plate 10, fig. 7, shows an ovocyte that has differentiated from 
the auxocyte; the first polar body (p. b^) is already expelled. 
The chromosomes are apparently getting into the diffused con- 
dition. Within the germinal vesicle are two "intra-nuclear 
division centers'' (i. dc), one of them apparently moving toward 
the periphery. Presumably this foreshadows the formation 
of the second polar body. A feature of the process is shown 
in the other ovocyte, where the attraction sphere or portion 
of it is now seen outside the germinal vesicle. Apparently 
the second meiosis, or formation of the second polar body, takes 
place by the flowing out of some nuclear materials into that 
portion of the attraction sphere outside the germinal vesicle. 
Evidence that such actually happens is the presence of two 
division centers in an older ovum (Plate 10, fig. 8, i. dc, e. dc) 
— one intra-nuclear, while the other is extra-nuclear. It seems 
that in this case the materials which had been withdrawn 
from the germinal vesicle supposedly to compose the materials 
of the second polar body were not cast off, but are disseminated 
within the ovum. Obviously, both maturation divisions in this 
group are reductional, as in S, tranquebarica. 

Plate 10 fig. 8, and Plate 11, fig. 1, show the young ova in 
different phases of growth. In the first, the nuclear frame- 
work, that is, the nuclear wall and the network, are still well 
defined. In the other figure, the network has disappeared; 

330 The Philippine Journal of Scienoe ^^® 

the germinal vesicle has increased considerably in size and so 
has the bulk of the cytosome. A spermatozoon (sp.) has pene- 
trated into the cytoplasm. 

Plate 11, fig. 2, is a photomicrograph of a large portion of a 
mature ovum* The yolk-spherule masses of various sizes ap- 
pear as blacfc Within the germinal vesicle, the sperm can be 
seen as a black mass. A line is drawn to mark the boundary 
of the ovum and that of the adjoining one. 


In this group, two classes of ovocytes are produced. As in 
S. tranquebarica, they are designated as the macro- and micro- 
ovocytes. In S. tranquebarica, however, the features of the 
changes that lead to the differentiation of these two classes of 
ovocytes from their respective auxocytes differ markedly. 

In the paramamosain the general nature of the changes is 
somewhat closely identical. The two classes of ovocytes dif- 
ferentiate from their respective auxocytes in about the same 
manner; in both, some kind of reorganization takes place in the 
nucleus, resulting in the sloughing off of some of the nuclear 
materials. Then, the remaining nuclear portion, together with 
a certain amount of cytosome, reorganizes to form the ovocyte. 
There are, however, features, that distinguish the two classes 
of ovocytes: (1) the marked difierence in size between the 
respective auxocytes including their nuclei; (2) the amount of 
cytosome involved, and the consequent difference in size of the 
ovocytes that develop, Plate 11, fig. 5, and Plate 12, fig. 3; (3) 
in the micro-ovocytes, the sloughed off materials form a mass, 
presumably forming the polar nucleus, Plate 12, fig. 3, p.n. 
This condition does not seem to be the case in the macro-ovocytes, 
wherein the sloughed off materials, apparently disseminate in 
the cytoplasm and presumably contribute to the materials of 
the ovocyte. Some of the characteristic features are shown in 
the following illustrations. 

Plate 11, fig. 3, shows a full-grown auxocyte (macro-class), 
apparently in the contraction figure or synisezis. In this case, 
the leptotene-threads do not seem to show any tendency towards 
definite polarization, while the chromatids appear to shift. A 
conspicuosuly large centrosome (c) with some centroplasm 
lies above the nucleus. 

Presunmbly the above condition is followed by the diffused 
stage, Plate 11, fig. 4. This is characterized by more or less 
diminished basophily and a loosening of the threads, which have 

^»» 3 Estampador: Studies on Spermatogenesis and Oogenesis 331 

become more or less deconcentrated. It seems significant that 
the prochromosome threads are still distinguishable and that 
they are distinctly double, the halves or the pairs being widely 
separated. The tendency towards polarization is shown in 
the nucleus, which is divisible into two zones, namely: (1) the 
portion adjoining the centrosome wherein the materials are 
completely diffused and (2) the other portion holding the pro- 
chromosomal threads which run towards the opposite pole. 
Obviously the noaterials within the completely diffused area 
are being disseminated into the cjrtoplasm. In other words, the 
nuclear elements are undergoing reorganization preparatory 
to the differentiation of the ovocyte. 

Plate 11, fig. 5, shows the second synizesis or contraction- 
figure. The nuclear substances are again concentrated in some 
degree. A similar change is also taking place in the cjrtoplasm. 
Apparently, the chromatids are being set in their alignment. 
One significant feature is the diminished size of the centrosome, 
compared with that in the preceding stages. The definitive 
form of the ovocyte is presumably attained at the completion 
of this phase. 

The differentiation of the micro-ovocjrte proceeds in a some- 
what similar fashion. Plate 12, fig. 1, shows a micro-auxocyte 
in the contraction figure, or synizesis. The nucleus shows dis- 
tinct polarization, the material in the region adjoining the cen- 
trosome are more diffused; these materials are apparently 
drawn to this zone, the threads which are rather rough in out- 
line are directed towards the centrosome. In the opposite pole, 
however, the threads are thinner and the chromatids are ap- 
parently being shifted. The nuclear figure may be regarded as 
similar to the so-called ''amphitene'' phase of some authors if 
polarization and the formation of two types of threads are taken 
as the sole bases. The case seems to be different, however, in 
the sense that the formation of the thicker threads is due to the 
crowding of nuclear materials preparatory to their being slouged 

Further contraction results in another contraction-figure, 
or synizesis, Plate 12, fig. 2. On the side adjoining the centro- 
some, a portion is pinched off from the main nuclear body. 
The shifting of the centrosome towards the apical region of 
the differentiating ovocyte seems to be associated with the some- 
what abortive nuclear division. Apparently the segregation 
of nuclear materials is taking place; the pro-chromosomal 

832 The Philippine Journal of Science ^^^ 

threads are shown radiating from the condensed, deeply stained 
mass located at the place of division. 

In Plate 12, fig. 3, the micro-ovocyte is shown to have com- 
pletely differentiated from the auxocyte body and the two nu- 
clear portions have also separated, the smaller one (p. n.) 
presumably representing the first polar nucleus. It appears to 
be within the centrosome. Wether the micro-ovocytes develop 
into functional ova is yet to be determined* There are indica- 
tions to the contrary. 

A photomicrograph (Plate 12, fig. 4) of a portion of the 
ovary shows that some of the developing ova appear to be stunt- 
ed (mi) ; presumably these are some of the micro-ovocytes. 
Some sperms (sp) may be seen outside the eggs; one (spz) is 
shown as having penetrated into the cytoplasm of a growing 
ovum and another (spO may be seen already inside the ger- 
minal vesicle. The heavy black ring that partially surrounds 
the germinal vesicle represents the vitellogenous layer, or pal- 
lial substance. It is apparent that in these crabs insemination 
may take place even before the ova become full-grown. 


The following illustrations are taken from the sections of the 
ovary of a young female, which measures 7.7 cm. across the 
carapace. The color of the ovary is orange, and from all ap- 
pearances, it looks normal like the ovaries in mature females. 
The cells appear to be developing normally. It is rather doubt- 
ful, however, whether the eggs can be fertilized in this stage; 
there are apparent obstructions in the form of the deep sternal 
concavity which partly hides the rather too small opening of 
the vulva. 

In this species, the process of odkinesis proceeds in a man- 
ner not very much different from that in S. oceanica. In both 
specie, the differentiation of the ovocyte takes place simul- 
taneously with the diminution of the nuclear materials and the 
formation of the first polar body. 

Plate 12, fig. 5, shows an auxocyte after the growth period. 
It presents a contraction-figure, or ssmizesis. Some of the 
threads radiate from the condensed heavy mass at the center, 
while those that are close to the border form loops. 

In Plate 12, fig. 6, the ovocyte has differentiated from the 
auxocyte iKHiy; the polar nucleus (p. n) and some nuclear 
materials have been extruded. The quadripartite constitution 
of the chromosome-nucleolus, which is presumably the X-chro- 

'*» 3 Estampador: Studies on Spermatogenesis and Oogenesis 333 

mosome, is distinctly shown. The prochromosomal threads are 
apparently undergoing realignment 

Plate 12, fig. 7, shows an ovocyte where concentration and 
contraction processes are apparently going on. The first polar 
body (p. n) is still embedded in the cytoplasm. The centrosome 
(c) with some centroplasm is distinctly visible. 

Plate 12, fig. 8, shows a contraction phase and the sloughing 
off of some cytoplasmic materials. The two polar bodies are 
still within the ovum. 

The form, size and character of the chromosomes are shown 
in Plate 12, figs. 9 and 10; the X-chromosome, or chromosome- 
nucleolus, is quadripartite. 


The principal points in oogenesis may be summarized as fol- 
lows : 

1. The auxocytes (ooblasts), after a period of growth charac- 
terized by a greatly increased bulk at the expense of the nurse 
cells, metamorphose directly into ovocytes. 

2. At the termination of the growth period, the auxocjrtes 
undergo transformation. The process is characterized by fea- 
tures that are quite distinctive for each group. 

3. In S. tranqueharica, odkinesis proceeds in two distinct 
ways: In one type, the differentiation of the ovocyte from the 
auxocyte involves practically the whole cell In the other the 
kinoplasmic area includes only the germinal vesicle and a very 
limited portion of the cytoplasm around it. In other words, 
the rest of the cell does not participate actively. Consequently, 
two classes of ovocytes are formed, namely: the macro-ovocytes 
and the micro-ovocytes. The macro-ovocytes complete the ma- 
turation period earlier; the formation of the second polar body 
follows inamediately that of the first polocyte, The ootid then 
proceeds to groW into mature ovum. In the micro-ovocyte the 
formation of the second polar body takes place when the ovocyte 
has become full-grown ovum, and after insemination has taken 
place. There is no corresponding ootid formation, and it seems 
that the penetration of the sperm into the germinal vesicle is 
necessary before the formation of the second polar body can 
take place. Throughout the period of growth to the time of 
the formation of the second polar body, the number of chromo- 
somes is six, but they are bivalent. The chromosomes become 
univalent after the formation of the second polar body. Ap- 
parently, the eggs of S. tranqueharica exhibit sexual dimegaly. 

334 TA^ Philippine Journal of Science ^^® 

4. In S. oceanica (banJiawin) the formation of the first and 
second polar bodies seems to run parallel with the differentia- 
tion of the ootid from the auxocyte. Only one class of eggs is 

5. The parmmamosain variety has more in common with S, 
tranquebarica in the sense that both groups produce two classes 
of ovocytes. One main difference between them lies in the fact 
that the micro-ovocytes in paraTnamosain do not appear to 
develop into normal eggs, in S. tranquebarica the two classes of 
ovocytes develop into functional ova. 

6. The process of ookinesis in S. serrata has much in common 
with that in S. oceanica; in both the differentiation of the ovocyte 
from the auxocyte takes place in conjunction with the formation 
of the polar bodies. Both produce only one class of eggs ; that 
is, they do not exhibit sexual dimegaly. 


During copulation the spermatophores are transferred from 
the vas deferens into the vulva of the female and into the sper- 
matheca where they are stored. By some means the spermato- 
phores find their way into the lumen of the ovarian tubules, 
where the spermatozoa are released from the enclosing capsules. 

If a crushed portion of a fr^h testis is examined under 
the microscope, tide material has the appearance of an alveolar 
substance containing numerous globules of various sizes. The 
bigger ones, which vary considerably in size, are probably fat 
globules. The spermatozoa app^r as very minute globular 
bodies resembling cocci bacteria, except that they show some 
sort of oscillatory movement. The tapering conical portion of 
the sperm body is not visible except in stained sections. 

The relation of the spermatozoa to the eggs taken from the 
lumen of the ovary of a parabanhawin (S. tranquebarica) is 
shown in Plate 8, fig. 11; Plate 9, fig. 3; Plate 10, figs. 1 and 
3, also in Plate 12, fig. 4, which is a photomicrograph of a 
portion of the ovary of a paramamosain. 

I cannot state definitely how the entrance of the sperm into 
the ova is effected in these crabs. Investigators on otiier crabs 
differ in their observations. KolimU (1906)^^ claimed that in 
certain decapods, the spermatozoa settle on the eggs with the 
clear cup towards the egg and the capsule pointed away from 
the egg. He believed that the rebound from the explosion of the 
capmile is sufiSk^ient to drive the nucleus into Ihie egg. 

Binford (1913)2 observed that in Menippe mercenaria the 
entrance ^f the sperm into the egg is effected by the turning 

^*^' 3 Estampador: Studies on Spermatogenesis and Oogenesis 335 

inside out of the tubule and capsule. According to him, the 
nucleus does not even enter the egg; hence, it does not partic- 
ipate in the process of fertilization. The fact that the nu- 
cleus holds the chromosomes regarded to be the bearer of pater- 
nal qualities to the egg^ and hence, are the essential thing in 
fertilization, he tried to reconcile by assuming that during the 
transformation of the spermatid, the nuclear material, at least 
the chromatin, is transferred to the capsule. He said: "the 
contents of the capsule may be derived from the nucleus of 
the spermatid and is probably oxychromatin which deposits 
basichromatin after it enters the egg and so gives rise to the 
chromosomes of the male pronucleus". 

In the present crabs the whole sperm enters the egg, and 
both nuclear cup and capsule play important rdle. But the 
mode of penetration and the dynamics involved are problems 
that need further study* It seems that some kind of an ever- 
sion which is explosive in nature takes place, whence the nuclear 
cup is turned inside out. This is suggested in Plate 8, fig. 11, 
and Plate 10, fig. 3. 

Insemination. — ^This process may take place as early as during 
the early part of the growth period of the ovum (Plate 8, fig. 11, 
and Plate 11, fig. 1) or later, when the egg has attained its fuM 
growth (Plate 9, fig. 3). The eggs are relatively so large and 
opaque and the spermatozoa so very minute that it is impossible 
to observe the actual process. Inference can merely be made 
only from a study of a series of sections of the fertilized eggs. 

By some means a sperm finds its way into the germinal vesicle, 
where the penetration path is discernible, but the path through 
the yolk-laden cytoplasm is entirely obscured. There is no 
actual bodily fusion between the male and the female pronu- 
clei; that is, they do not unite to form a single nucleus, or con- 
jugation nucleus. 


The following description is based upon the eggs of S. tran- 
quebarica (parabanhawin) . 

The enormous increase in the size of the ovum is accompanied 
by that of its germinal vesicle. Two classes of ovocytes pre- 
viously referred to as macro- and micro-ovocyte were observed 
in parabanhawin. The germinal vesicle in the micro-ovocyte 
is enormously big, compared with that in the macro-ovocyte 
(Plate 10, figs. 1 and 3). The general framework of the ger- 
minal vesicle of the micro-ovocyte is distinctly oxyphilic; the 

336 T^ Philippine Journal of Science *^^ 

framework constitutes the major part and there is a decided 
tendency for the chromatin materials to collect at the periphery 
(Plate 10, fig, 2.) 

After the entrance of the sperm, the germinal vesicle begins 
to fragment (Plate 10, figs. 3, 4) ; the karyoplasm becomes 
organized into karyomerites (k) and karyomicrosomes (m), or 
nuclear microsomes. The former are masses of karyoplasm con- 
taining chromosomal elements, while the latter are only granu* 
lar masses or karyoplasm. These masses stream out of the 
germinal vesicle (Plate 10, fig. 4) ; hence, the germinal vesicle 
gradually becomes smaller and smaller and finally disappears 

The tendency of the chromatin materials to align at the peri- 
phery of the germinal vesicle (Plate 10, fig. 2) assumes added 
significance if coupled with the apparent polarity of the forma- 
tion of karyomerites (Plate 10, fig. 3, fc). Presumably these 
phenomena are associated with the formation of the second polar 
body and may be regarded as phases leading to the segregation 
of the potential functional pronuclear elements from those which 
are to constitute the materials of the second polar nucleus. 
The following events seem to support this idea: 

At about the time the germinal vesicle is reduced to about 
half its original bulk, resulting from the extrusion of the kary- 
omerites and nuclear microsomes, the remaining part begins to 
differentiate into two portions. One of these portions event- 
ually segregates and becomes the second polar body. The 
framework of this portion is deeply oxyphilic and appears more 
homogeneous than the other portion. In this latter, there are 
indications that the process of fragmentation is proceeding, be- 
cause its wall is indistinct; the ground substance merges with 
the outlying materials which are largely composed of minute 
masses similar to those present within this body and in xne 
karyoplasmic streamers. Eventually this portion disappears 
as a body, leaving in its place a homogenous granular mass. 
A number of masses similar to this is scattered in various 
parts of the egg. They are obviously cytoplasmic masses to- 
gether with some nuclear materials (karyo-cytoplasmic masses.) 

The formation of the second polar body in tiiis type of eggs 
(micro-ova) may be summarized as follows : The potentioal for- 
mative pronuclear elements are sorted out in the germinal 
vesicle through the operation of agencies which are not known. 
The entrance of the sperm into the germinal vesicle initiates 

^*' 3 Estampador: Studies on Spermatogenesis and Oogenesis 337 

the movement of the karyomerites and karyo-microsomes out 
from the germinal vesicle, and when the latter is reduced to 
about half of its original bulk, it splits into two portions. One 
of them becomes the second polar body, the extrusion of which 
from the egg takes place presumably during cleavage. 

The yolk-spheres (Plate 9, fig. 3) appear at first as more or 
less homogenous masses, conunonly bearing some pigments. 
The homogenous aspect of the spheres is transformed later to 
that of a vacuolated appearance, becoming divided into separate 
masses of deeply staining materials and interspersed by non- 
staining areas. They soon break up into smaller vacuolated 
bodies, which then gradually break up into small granular 
masses. Finally, the ooplasmic framework in the egg appears 
to be divided into somewhat homogenous blocks, separated by 
faintly staining to nonstaining lines. These lines presumably 
represent the protoplasmic strands that radiate from the center 
of the egg to the superficial cytoplasmic layer, the blastema. 
It is along these somewhat clear spaces that active formation of 
protoplasmic islands takes place during cleavage and, con- 
sequently, where the active assimilation of the yolk is going on. 
Along some of these areas are formed the secondary cleavage 
of some authors. 


When a crab is ready to spawn, she raises her body and 
assumes an upright position holding the abdomen away from 
the body; the abdomen together with the exopods form a sort 
of a basket into which the eggs are run. The eggs become at- 
tached to the setae of the endopodites, where they pass their 
embryonic stages. 

According to the observation of Mr. Arriola (1940), these 
crabs spawn throughout the year, the peak of activity falling 
between May and September. This observation was made at 
the Experimental Station of the Division of Fisheries at Dagat- 
dagatan, Malabon, Rizal Province. My observations on the 
conditions of the gonads of crabs dissected during different 
months of the year corroborated Arriola's findings. 

That the period of the peak of spawning activity may vary 
perhaps slightly in different regions of the Philippines is prob- 
able. For example, in Dumangas, a town in Iloilo Province 
noted for the abundance of alimango, the season of highest 

338 The Philippine Journal of Science *^* 

spawning activity appears to be between August and Novem- 
ber. During the intervening period, crabs with gonads full of 
eggs are sold abundantly in the markets. 

Mr. Arriola (1940, p. 443)^® presumed that the crabs die 
shortly after laying one batch of eg^. The presumption is 
based on the behavior of his specimens, which were kept in 
floating cages. I can not entirely subscribe to his view. In 
the first place, it should be borne in mind that the crabs are dis- 
tinctly bottom-dwellers and fo^orial in habit. The mamosain, 
for instance, live in burrows, while the others live concealed in 
mud-holes and under shelter among heaps of debris. By placing 
the animals in floating cages, he subjected them to unnatural 
conditions. Certainly, it is not safe to draw conclusions pur- 
porting to be the same with what is expected to happen under 
natural conditions, especially when the experiments were per- 
formed under conditions at variance with the natural setup 
wherein the animal thrives favorably. 

In tiie second place, his assertion does not seem to find sup- 
port from facte observed under natural conditions. According 
to his observations, these crabs attein sexual maturity when 
they reach the siaie of about 84 by 122.5 mm. across the cara- 
pace. Possibly, his specimen must have been a true banhawin, 
for I had in my collection a berried parabcmhawin that measures 
only 100 mm. across the carapace. From the studies on the 
sections of ovaries and testes from crabs of different sizes, 
I found female crabs of 90 mm. across tiie carapace already 
with mature gonads. Nevertheless, just for the sake of having 
sometiiing to base on, let us teke 84 by 122.5 mm. as the size 
at which some of these crabs attein sexual maturity. What 
can we say of some female crabs reaching the size of 200 mm. 
or 225 mm. across the carapace? Big crabs of tiiese sizes are 
not very rare. Certeinly, we cannot say that thaie crabs had 
not spawned previously. 

Egg menbrane, — ^Upon reaching the exterior, each egg is sur- 
rounded by two manbranes: the vitelline (or fertilization) mem- 
brane and tiiie chorion. Between these two mambran^ is a 
perivitelline space which, according to Williamson, ^^ a>nteins 
a fluid possessing adhessive properties. Microscopic studies 
made on stained i^ctions of both ovarian eggs and on eggs 
already attach^ to the endoiK^ite setse did not rev^l any 
indication of tihie presence of this fluid. On the other hand, the 
two membranes are intimately fused in some plac^, though 

W' 3 Estampador: Studies on Spermatogenesis and Oogenesis 339 

generally the two membranes show separately (Plate 13, fiir. 
landPlatel4, fig. 1). 

How the eggs of crabs become attached to the endopodite 
setae is a question that has not been satisfactorily answered. 
Some of the earlier observers believed that a sticky substance 
is secreted around the eggs as they are shed. The objection 
to this theory is that it does not explain why the eggs become 
attached only to the endopodite setse. The view given by Her- 
rick (1895) is that the tegumentary glands of the endopodites of 
the pleopods secrete an adhesive fluid. However, there are 
physical aspects which this theory implies that cannot be ex- 
plained. In the first place, if the eggs became attached as they 
came in contact with the surface that secreted this adhesive 
fluid, then some of the eggs should adhere to the endopodite 
itself. This does not seem to happen, however. In the second 
place, there are possibilities for the eggs to become attached 
together in groups around a point. This does not seem to hap- 
pen either. There are other aspects that point to the unaccept- 
ability of the premise. 

A different theory is suggested by Williamson. He explains 
that the endopodite setse penetrate the chorion of the egg in 
two places, and thus the egg becomes skewered on the seta. 
The piercing of the chorion liberates the adhesive perivitelline 
fluid which assists in making the attachment more permanent, 
and the egg appears to be attached to the seta by a stalk. 

Williamson's theory implies two things: (a) either there must 
be some kind of a guiding mechanism that enables the seta 
to pierce the egg in two places or (b) this piercing takes place 
merely by chance. Since the setae are very fine and long, they 
must be in motion constantly, moved by the water currents 
which must be especially active at the time the eggs are being ex- 
truded. The extrusion of the egg itself will cause some water 
movements. No such mechanism implied in the theory can 
operate under such conditions. If the piercing of the chorion 
is just a matter of chance, then the probability is that many of 
the eggs will be lost, if not the majority of them. Sometimes 
the number of eggs attached to a seta reaches a hundred or more; 
the eggs are ranged from the base to the tip of a seta. This 
condition cannot be possible with the mechanism implied in the 
theory. Moreover, Williamson's idea about the presence of an 
adhesive perivitelline fluid is not supported by the findings from 
cytological studies. 

9543 5 

340 The Phil^f^pine Jourtud of Science ^^^ 

The most plausible explanation seems to be that a sticky 
substance is secreted around the eggs as they are shed. To 
the question as to why the eggs do not become attached to the 
exopodite setae, the following answers may be given: 

(1) The topographical position of the exopodite setae, their 
being on the outside, reduces the chances for the eggs to be- 
come attached to them. 

(2) The eggs on being released outside, first come in con- 
tact with the endopodite setae, which are so thick that th*ey 
form a sort of an impenetrable barrier. Their tangled mass 
affords the eggs all chances of becoming attached, and at the 
same time help keep them within the confines of the entangle- 

(3) The structural make-up of the exopodite setae and their 
peculiar arrangement render them less adapted for the attach- 
ment of the eggs. They are shorter and finer, and possess side 
branches (Plate 1, fig. 5, 6). These branches interlock forming 
a more or less compact surface, and together with the broad 
expanded surfaces of the exopodite stalks, a sort of a wall is 
formed, guarding the sidespaces between the sternal wall and 
that of the abdomen, when the latter is drawn away from the 
cephalothorax during the act of spawning. In other words, 
a sort of basket is formed with the abdomen serving as the 
bottom and the exopodite stalks and their setae form the wall. 
One has only to see a berried crab in order to form an idea, that 
the exopodites and their setae are structural devices adapted to 
act as barriers. 

There are other factors that prevent the eggs from attaching 
to the exopodite setae, namely: (1) the mobility of the exopo- 
dites due to their being loosely joined at the basal segment 
and (2) the water current. The current set by the movement 
of the water as it is drawn into the inhalant chamber glides 
down the sternal wall ; its impact upon the broad expanded sur- 
faces of the exopodites spreads the latter outward. The eggs 
as they glide down within the sternal concavity also produce 
some water movement. Cross-currents are then produced with- 
in the stemo-abdominal space. The interplay of the currents 
helps to prevent the eggs from attaching to the exopodite setae 
and at the same time keeps them within the confine of the 
sterno-abdominal space amongst the endopodite setae. The 
water current also keeps the eggs apart from one another and 
sets them in motion amongst the setae until they become at 

'*» ^ Estampador: Studies on Spermatogenesis and Oogenesis 341 

tached. It is surprising to note that the eggs do not become at- 
tached to one another. 

The adhesive fluid by means of which the eggs become attach- 
ed to the endopodite setae is produced by the secretory cells 
(Plate 7, fig. 9, c, c) of the ovarian follicles. This substance 
was previously referred to as something collagenous in character. 
While on their way to the outside, the eggs become coated with 
this substance which enables them to adhere to the endopodite 
setae with which they come in contact. The stalk with which an 
egg appears to hang on a seta presumably is formed in this 
manner: the weight of the egg stretches out a portion of this 
sticky substance and in the meanwhile the action of the water 
gradually hardens it. Microscopic studies on stained sections 
of the eggs reveal the presence of a layer of this material 
around the chorion, and this layer is continuous with the ma- 
terial that forms the stalk (Plate 14, fig. 1, st). 


The development may be divided into three stages, namely 
embryonic, larval and postlarval. 

The embryonic development takes place while the egg is at- 
tached to the pleopods of the mother. Since the embryonic 
stage has not been observed in the present study, an attempt 
will merely be made to give a fragmentary sketch. 

Cleavage. — The entrance of the sperm into the germinal ves- 
icle initiates radical changes in this body. It begins to frag- 
ment into karyomerites and karyomicrosomes (Plate 10, fig. 3). 
In the meantime the sperm body also begins to undergo reor- 
ganization inside the germinal vesicle. It becomes vesicular 
and then begins to fragment into karyomerites (Plate 10, figs. 
l,sp). In this fragmentation process, the idiosome (attraction- 
sphere) seems to play a very important role. It should be re- 
called in this connection that the idiosome and the acrosome 
materials form the capsule of the sperm. The fragments of 
the sperm together with those of the germinal vesicle stream out 
and this process continues until the germinal vesicle disappears 
as a body. 

The fragments from both the sperm and the germinal vesicle 
find their way into the cytoplasmic areas scattered amongst the 
yolk masses. Presumably it is within these cytoplasmic areas 
where the paternal and maternal elements come into intimate 
association and where the male and the female pronuclear ele- 
ments do actually unite. 

342 2^*^ Philippine Journal of Science *•** 

It is then apparent that syngamy does not take place by the 
direct bodily fusion of the egg and sperm nuclei ; it is effected 
in a somewhat diffused manner by the fusion of their essential 
fractional units (karyomerites) in the karyo-cytoplasmic areas. 
Each ssygote-karyomerite resulting from this fractional syngamy 
seems to influence the active changes that take place around 
its immediate vicinity. 

Starting from those located near the center of the egg, pro- 
found changes then take place in the karyo-cytoplasmic areas. 
They soon begin to form into more or less separate irregular 
masses, into a number of nonstaining areas, each surrounded by 
protoplasm; these are the so-called "protoplasmic islands". 
Some authors consider the protoplasmic islands as cleavage cells, 
although they are not cells in the sense compared to other types 
of segmenting eggs. By continuous division, they approach the 
periphery of the egg and finally unite with the superficial cyto- 
plasmic layer (the blastema) . Then the cytoplasmic boundaries 
become visible and the layer becomes a blastodermic epithelium 
— ^the layer assuming the appearance of being divided into plas- 
ma blocks (Plate 13, fig. 1). Since the divisions do not extend 
through the yolk, cleavage in this crab is distinctly superficial. 

In each plasma block there is generally present in stained sec- 
tions a big, heavily stained mass. The presence of this peculiar 
structure indicates that the blocks are really syncytial masses 
of cytoplasm and chromatin materials, and possibly including 
some pigments and other ooplasmic substances. The absence of 
protoplasmic islands in the vicinity of the heavily stained mass, 
in sharp contrast to the presence of several of them in blocks 
that do not contain darkly stained masses, seemingly supports 
this view. 

The term "protoplasmic island" in this sense merely purports 
the presence of nuclear materials in a mass of cytoplasm, rather 
than the existence of fully formed nucleus, since its formation is 
traceable to the inflow of karyomerites of both the sperm 
nucleus and those of the germinal vesicle into the cytoplasmic 
area amongst the yolk-masses. The view will hold, if we con- 
ceive the idea that these nuclear fragments become reorganized 
into full-fledged nucleus within the cytoplasmic mass. This idea, 
however, recognizes total potentiality of a portion (the karyo- 
merite) to reproduce the whole chromosomal complements of 
a cell. This contravenes the established idea regarding the 
individuality of chromosomes since a karyomerite is only a frac- 
tional portion of a nucleus. 

^^ ' Estampador: Stvdieson Spermatogenesis and Oogenesis 343 

Presumably, what takes place in a cytoplasmic mass is that 
a karyomerite from the sperm nucleus unites with one from the 
germinal vesicle of the ovum to form a zygote-karyomerite. 
This zygote-karyomerite soon undergoes a series of divisions 
forming a number of daughter karyomerites which then undergo 
reorganization and become what we may regard as ''protoplas- 
mic islands." The formation of plasma blocks may be inter- 
preted as a phase tending toward bringing together what had 
been scattered before through nuclear fragmentation. In other 
words, the chromosomal complement that each karyomerite is 
supposed to carry must be brought together with those borne 
by the other sister karyomerite before the organization of the 
nucleus of a cellular unit can proceed; that is, in some sort of 
a reassociation. The plasma block was previously referred to 
as a syncytial mass of cytoplasm, chromatin elements, and pos- 
sibly with some ooplasmic and other materials. The formation 
of a number of cells from each block in the later stages of 
development seems to bear out this inference — that the reorga- 
nization taking place within the blocks is a counterphase leading 
to bringing about nuclear reconstruction, and foreshadowing the 
formation of definitive cellular units. 

Some investigators regard the nuclei (karyomerites in this 
case) that remained in the yolk as "vittelophags," because they 
assume the special function of the assimilation of the yolk. 
Around these vittelophags some incomplete boundaries are 
formed giving rise to what may be called secondary yolk- 
cleavage. The vittelophags are regarded by some authors as 
endoderm, and some investigators have described superficial 
cleavage as leading directly to the formation of a gastrula-like 
stage, since two kinds of cells, ectodermal and endodermal, 
develop about the same time. 

Some authorities classify superficial cleavage into two groups : 
(1) cleavage at first total but later superficial; (2) cleavage 
purely superficial. Both groups may again be divided into two 
subdivisions, namely: (a) the blastomeres forming on all sides 
simultaneously and (b) the blastoderm in the ventral side 
developing precociously. 

Cleavage in these crabs (S. tranquebarica) is purely super- 
ficial, and may be classed under the second subdivision ; that is, 
the blastoderm in the ventral region develops precociously (Plate 
13, fig. 2). The character of the cleavage is determinative; 
that is, the cjrtoplasm of the egg and the regions are clearly 
organized in relation to their future destiny. 

344 The Philippine Journal of Science ^^* 

The egg is ovate in outline, the narrower end being the future 
anterior part of the body. There seems to be more cytoplasmic 
masses in the presumptive ventral side, but fewer yolk-masses ; 
whereas, in the presumptive dorsal side the yolk-masses appear 
to be more numerous. Perhaps the precocity in the develop- 
ment of the ventral side can be attributed to the presence of more 
cytoplasmic masses, in which region the process of development 
is less hinderd by the yolk material. 


The movements of protoplasm are quite discernible and at 
least from the mechanical point of view they may throw some 
light on certain features of development. 

The ovarian egg has its cytoplasm in a delicate superficial 
layer (blastema) and in numerous delicate intravitellar strands 
connecting this with the cytoplasmic layer around the adcentral 
germinal vesicle. In some places among the yolk-spheres, the 
cytoplasm collects in masses forming what we may term "cyto- 
plasmic areas." These partake the nature of outposts where 
the karyomerites of the male and the female pronuclei presum- 
ably meet and unite. After fertilization these areas become 
"karyo-cytoplasmic" masses ; starting from those located at the 
center, these masses soon undergo division, giving rise to nume- 
rous protoplasmic islands. 

The cleavage of the eggs consists of movements leading to 
the transposition of both nuclear and cytoplasmic materials to 
the surface. This is in a sort of general cytoplasmic movements 
of the nature of either a flowing or a contraction. The latter 
movements are in part upon the yolk surface ; in part they consist 
of movements of vitellocytes within the yolk, but they are 
blastodermic and not vitellar. 

Cleavage in this crab does not lead to a definite blastula and 
gastrula formation. These two stages, ordinarily occuring as 
distinct phases, are indistinguishable in this case. 

The early blastoderm consists of a peripheral layer of plasma 
blocks (Plate 13, fig. 1). The ventral germ-disc becomes estab- 
lished by more rapid formation of plasma blocks, and to a cer- 
tain extent by the apparent rapid migration of protoplasmic 
islands towards this side. Cell membranes begin to appear while 
the germ disc is forming; prior to this time the vitelline mem- 
brane and the cytoplasm represented one great reticular syncy- 

^^» * Estampador: Studies on Spermatogemsis and Oogenesis 345 

tium, composed of more or less definite masses, the "plasma 

In the absence of definite cell wall and of definite nuclear 
body, the term "plasma block" is quite appropriate for these 
masses. They soon, however, give rise to cells with distinct 
membranes. While this is taking place around the periphery, 
plasma blocks are also formed within the embryo. The rate of 
proliferation in the peripheral plasma blocks differ; consequent- 
ly, the embryo appears to be divided into independent territories 
wherein active changes are taking place. 

Growth differences between cephalothoraz and abdomen. — The 
cephalothorax and abdomen show certain striking differences 
in their growth and differentiation. The boundary between the 
two can be recognized as a thin region of the ectoblast where 
a certain degree of insinking takes place and active proliferation 
of the cells in the adjoining regions (Plate 14, fig. 1) appears 
to take place. 

Nearly the whole extent of the cephalothorax is laid down, 
thereafter to lengthen but little more, when the abdomen is 
represented only by a small lobe (Plate 14, fig. 1, c. I). With 
the segmentation of the mesoblast the abdomen commences to 
increase rapidly in length, pushing around the yolk until the 
caudal lobe finally comes in contact with the head lobe. Its 
growth is teloblastic, and the successive segments formed from 
its anterior end caudal. This rapid elongation of the abdomen 
explains why the caudal lobe is narrower than the cephalic lobe. 
Segmentation becomes more distinct in the abdomen (Plate 14, 
fig. 2, ab). 

Protozoea (Plate 14, fig. 3). — Hatching takes place at this 
stage. The large paired eyes are present; there are no frontal 
or dorsal spines. The abdomen is well defined, but only five 
somites and the telson are distinguishable. The sixth somite 
is still fused with the telson; the latter forked with strong 
spines in each branch. All the cephalic and the first two pairs 
of thoracic limbs are present. The antennule consists of a basal 
segment from which arise two branches and each fork is armed 
with rather long setae. Antenna consists of two segments 
with a few terminal setse. It is postoral in position. Ap- 
parently during the stage of reversion it is shifted anterior to 
the mouth. The mandible is a small plate. The first and second 
maxillse are similar. The first and second pairs of maxillipeds 
are large and biramous. The protopodite is large and the en- 

346 The Philippine Jourmd of Science ^•*® 

dopodite and the exopodite have few sete* As shown in the 
illustration, the cuticular covering is becoming loose; apparently 
this indicates that the larva is about to change into the zoea 


1. Nichols, Louise M. Comparatiye studies in crustacean spermatoge- 

nesis. Jour, Morph. 20 (1909) 461-478. 

2. BiNFORD, Raybhjnd. The germ cells and the process of fertilization 

in the crab, Menippe mereena/ria. Jour. Morph. 24 (1913) 147-200. 

3. Pearson, Joseph. Cancer. Liverpool, Marine Biological Committee, 

No. 16 (1908) 1-207. 

4. Wilson, Edmund B. The cell in development and heredity (1928). 

5. DoNCASTBB, Leonard. An introduction to the study of cytology (1924). 

6. Darlington, C. D. Recent studies in cytology (1937). 

7. Richards, Aute. Outline of comparative embryology (1931). 

8. Huxley, T. H. The crayfish (1906). 

9. Hbrrick, F. H. The american lobster. Bull. U. S. Fish Comm. (1895). 

10. Arriola, F. J. A preliminary study of the life history of Scylla serrata 

(Forskal). Philip. Jour. Sci. 73 (1940) 437-454. 

11. Williamson, H. C. Twenty-third report, Scotch Fishery Board. 

12. KoLTZOFF, N. K. Studien iiber die Spermien der Decapoden, als Ein- 

leitung das Problem der Zellengestalt. Arch, f, ndkros. Anat. 
Bd. i7 (1906). 


PlJlTB 1 

Fig. 1. Ovigerous female, with chelipeds removed, p. o, genital opening; 
i o, inhalant opening. 

2. Ovaries, X 1. c, chitinous portion of the spermatheca; g, grandular 

portion of the spermatheca; g, nit gonadial memhrane; od, 
oviduct; ov, posterior extension of the ovaries; ov, /, ovarian 
follicle; spt, spermatheca; v, vulva. 

3. Cross-section of ovary through region above chileped, X 2. bv, 

blood vessel; c, canal formed by the ligamental fold; d.w, dorsal 
wing; t, one of the follicular tubules; v.Wf ventral wing. 

4. Abdomen of female 5. tranquebarica, showing character of seg- 

mentation, X 1. 

5. Second abdominal appendage, of a female, X 2, a, coxopodite; 6, 

basipodite; en, endopodite; ex, exopodite; s, fringe of setae. 

6. Portion of exopodite seta, showing dendritic characteristic, X 600. 

Plate 2 

Fig. 1. Abdomen of male S. tranquebarica, showing character of seg- 
ments, X 1. 

2. First abdominal appendage of male, modified as copulatory ap- 

paratus; left side, inner surface, X 2. a, basal joint; fr, groove; 
c, cistema spermatica; d, excavated area; e, groove along fun- 
nel tube. 

3. First abdominal appendage of male, modified as copulatory ap- 

paratus; right side, outer surface, X 4. a, tip of the funnel 
tube; by free edge of rolled portion of tube; d, canal leading 
to mouth of tube. 

4. Second abdominal appendage, also modified as copulatory appa- 

ratus. Right side, dorso-lateral surface; enlarged, c, canal 
in the fork; /, fork; w/, membranous fold. 

5. Second abdominal appendage, terminal portion only, inner surface 

to show the spinules on trough of fork, enlarged. 

6. Testes XI. ed, ajaculatory duct; /, follicular tubule; gm, gonadial 

membrane; ps, pedal sinus; U, testis; vd, vas deferens. 

7. Two epithelial cells from lining of lumen of ejaculatory duct, 

X 1350, camera lucida. a, attachment; b, eo, basal corpuscle 
to each cilium; e, cilia; c6, ceU body; /, fibrils corresponding 
to each cilium; n, neck. 

Plate 3 

Fig. 1. A growing auxocyte in follicular tubule of testis of S. tranqueba- 
rica, X 1350, camera lucida. 
2. A pro-spermatid undergoing reorganization. The spiremes radiat- 
ing from heterochromosomes, the darkly stained mass at center 
of germinal vesicle, X 1350, camera lucida. 


348 'The Philippine Journal of Science *^® 

3. A pro-spermatid undergoing further reorganization. The spiremes 

are transformed into chromatids and their groupings indicate 
tetrad formation, X 1350, camera lucida, l, a pair of tetrads. 

4. Pre-synaptic stage, showing number of chromosomes and pairing, 

X 1350, camera lucida. K heterochromosomes (the XY-pair) ; 
X-chromosome in form of loop, Y-chromosome small; m, mono- 

5. Chromosomes in synaptic stage; appearance of the idiosome and 

acroblasts, X 1350, camera lucida. h, heterochromosomes ; i, idio- 
some; m, monosome; p, ac, pro-acrosome granules. 

6. Pro-spermatid body in process of reorganization, showing concen- 

tration, contraction and sloughing-off of materials, X 1350, 
camera lucida. ac, acroblast (central body); g, Golgi-bodies 
remnant; i, idiosome; ne, nuclear extrusion. 

7. Pro-spermatid undergoing concentration and contraction; X 1350, 

camera lucida. i, idiosome; m, monosome, the extra X-chromo- 
some. The chromosomes are now dyads and the acroblast body 
has disaggregated. 

8. Formation of definitive spermatid body; sloughing-off of residual 

protoplasm and diminution of both nuclear and acroblast mate- 
rials, X 1350, camera lucida. c.e, possibly cast-off nuclear 
or acroblast materials; i, idiosome; r,p, residual protoplasm; 
8p,b, spermatid body; a?, sex-chromosome. 

9. Spermatid undergoing abortive mitotic division involving largely 

the nucleus; this results in neiosis, X 1350, camera lucida. 
ac, acroblast; d, daughter nucleus; m, probably the extra X- 
chromosome ( monosome ) . 

10. Top view of spermatid, wherein process of concentration is going 

on, X 1350, camera lucida. ac, acroblast; i, idiosome. 

11. Side view of spermatid in contraction phase, X 1350, camera 

lucida. ac, acroblast; r.p, residual protoplasm. 
12. Basal-side view of spermatid undergoing concentration, X 1350, 

camera lucida. acr, acroblast-remnant. 
13. Basal portion of spermatid, showing number of chromosomes, X 

1350, camera lucida. 
14-18. Later phases of transformation characterized by peculiar behavior 

and movement of acrosome body, X 1350, camera lucida. 
19. Spermatozoon, X 1350, camera lucida. ca, capsule; nc, nuclear 


Plate 4 

Fig. 1. Developing auxocyte (of class without tetrads), drawn from 
cross-section of follicular tubule, X 1350, camera lucida. 
/, follicular wall; ch.n, chromatin nucleolus;, presumably 
nucleus of nurse cell. 

2. Surface view of auxocyte (without tetrads), X 1350, camera lu- 

cida. nx, nurse cell. 

3. Side-view of auxocyte (without tetrads), at termination of growth 

period, X 1350, camera lucida. /, follicular wall. 

4. Chromosome pairs of auxocytes without tetrad formation, X 1350, 

camera lucida. h, heterochromosomes (possibly XY-pair). 

^»'^ Illustrations 349 

5. Showing two spermatida that have differentiated from auxocytes 
(of class with no tetrad formation), X 1350, camera lucida. 
acy acroblast; g, Golgi-bodies remnant; n.c, nurse cell; ti.e, 
nuclear extrusion; sp, spermatid body; v,ac, acroblast vesicle. 
6-10. Later phases of transformation of spermatids (of class with no 
tetrad formation), X 1350, camera lucida. ac^ acroblast; axi,r, 
acroblast-remnant; ca^ capsule; t, idiosome; n,c, nuclear cup. 

Plate 5 

Fig. 1. Group formation of auxocytes (spermatocyst), X 1350, camera 
lucida. cySf cyst; sp, spermatoblast (auxocyte). 

2. Auxocyte from group formation (spermatocyst), showing chromo- 

somes, X 1350, camera lucida. Y-chromosomes {y) is small, 
apparently triosomic and attached to big X-chromosome. 

3. Auxocyte of S, oceanica, in which nucleus has split into two sper- 

matid nuclei, X 1350, camera lucida. a, attraction sphere; c, 
centriole; /, follicular wall; sp. n, spermatid nucleus. 

4. Spermatid that has developed from spermatid nucleus, X 1350, 

camera lucida. 
5-7. Some phases of transformation of spermatid (in Sf. oceanica) to 
spermatozoa, X 1350, camera lucida. ac, acroblast; acr, acro- 
blast remnant; i, idiosome. 
8. Spermatozoa (S. oceanica), X 1350, camera lucida. a, Basal- 
side view; b, Basal portion only. 
9-10. Chromosomal complement (S, oceanica), X 1350, camera lucida. 
11. Pro-spermatid of paramamosain, showing nuclear diminution and 
slougking-off of some cytoplasmic material, X 1350, camera 

Plate 6 

Fig. 1. Somewhat more abbreviated process of pro-spermatid differentia- 
tion from auxocyte (in paramamosain), X 1350, camera lucida. 
2. Formation of two spermatids from pro-spermatid body, X 1350, 
camera lucida. 
3-4. Some phases of transformation leading to formation of sperma- 
tozoa, X 1350, camera lucida. 

5. Spermatozoa, X 1350, camera lucida. 

6. Photomicrograph showing spermatocyst of S. serrata, X 400. 

Note that some spermatids appear in pairs. 
7-8. Process of reorganization in nucleus of auxocyte (of S, serrata) 
leading to differentiation of spermatid, X 1350, camera lucida. 
c, centrosome with some centroplasm. 
9. Differentiation of spermatid nucleus from auxocyte nucleus (in S. 
serrata), and sloughing-off of some material, X 1350, camera 
lucida. r. p, sloughing-off material. 
10-12. Phases of nuclear reorganization in auxocyte nucleus (in S. ser- 
rata), tending toward differentiation into two spermatid nuclei, 
X 1350, camera lucida. 

350 The Philippine Journal of Science *^^ 

Plate 7 

Fig. 1. Phases of nuclear reorganization in auxocyte nucleus (in S, ser^ 
rata) J tending toward differentiation into two spermatid nuclei, 
X 1350, camera lucida. 

2. Nucleus, with chromosome-nucleolus at equatorial region prior to 

its splitting, X 1350, camera lucida. 

3. Division, X 1350, camera lucida. cy, portion of cytoplasm of 

auxocyte; i, idiosome. 

4. Side view of remains of auxocjrte; the spermatids are still within, 

X 1350, camera lucida. 
5~7. Later phases of spermatid transformations, X 1350, camera lucida. 

8. The spermatozoon, X 1350, camera lucida. 

9. Cross-section of ovarian follicle, X 600, camera lucida. c, some 

collagenous material in lumen of follicle; ex, collagen-forming 
cells; /, follicular wall; ^, lumen of follicle; w, nurse cell; o, 
growing ooblast (auxocyte). 

10. Two developing auxocytes together with some nurse cells, from 

section of surface of ovarian tubule of S, tranquebarica, 
X 1350, camera lucida. 

11. Developing auxocyte together with some nurse cells, from cross- 

section of ovarian tubule of S. tranquebarica, X 1350, camera 
lucida. /, follicular wall; n. c, nurse cell; a, developing ooWast 

Plate 8 

Fig. 1. Auxocyte at termination of growth period, X 1350, camera 
lucida. en, chromatin-nucleolus; g, Golgi-bodies; n, germinal 
vesicle; nc^ vestiges nurse cells; v, vitellogenous or pallial sub- 

2. Ovocyte in pachytene stage of contraction phase, X 1360, camera 

lucida. Irregularly scattered spherules appear in place of pal- 
lial substance. 

3. Diploid number of chromosomes, supernumerary chromosomes and 

diminution of nuclear materials, X 1350, camera lucida. h, 
hetero-chromosomes, evidently XX-pair. 

4. Ovocyte, showing extrusion of first polar body, X 1350, camera 

lucida. p.6i, first polar body. 

5. Ovocyte apparently in contraction phase, after formation of first 

polar body, X 1350, camera lucida. 

6. Pairing of chromatid groups prior to meiotic division which in- 

volves nucleus only, X 1350, camera lucida. h, hetero-chromo- 
somes; m, monosome; p, pb, path of first polar body. 

7. Chromosomes in secondary pairing prior to meiosis, X 1350, 

camera lucida. h, hetero-chromosomes; a.s, attraction sphere 
with two division centers; m, monosome. 

8. Meiotic division involving germinal vesicle only, X 1350, camera 

lucida. This will result in formation of second polar body. 

9. Ovocyte with germinal vesicle divided into two daughter nuclei, 

X 1350, camera lucida. p. &2, the second polar body. 
10. Young ovum with second polar body still within its body, X 1350, 
camera lucida. The egg pronucleus is turning around towards 

^*' ^ lUttstrationa 351 

11. Young ovum showing: penetration of sperm into ooplasm, X 1350, 
camera lucida. ch, chorion; m, path of penetration; ps, peri- 
vitelline space; sp, sperm body; v, vitelline membrane. 

Plate 9 

Fig. 1. Ovocyte (of class with no tetrad formation) differentiating from 

auxocyte body, X 1350, camera lucida. g, Golgi-bodies ;, 
first polar body. 

2. Extrusion of first polar body, X 1350, camera lucida. Both 

ovocyte and polar nucleus still within auxocyte body, a, attach- 
ment of the auxocyte to the follicular wall; h, hetero-chromo- 
somes; p,n, polar nucleus; o, ovocyte. 

3. Fertilized ovarian ovum (macro-ovum), X 900, camera lucida 

6, peripheral cytoplasmic layer (blastema); ch, chorion; cy, 
cytoplasmic mass; g.v, germinal vesicle; sp, sperm vesicle; y, 

Plate 10 

Fig. 1. (terminal vesicle of macro-ovum, with sperm inside, X 1350, camera 
lucida. Sperm evidently undergoing reorganization, cy, 
cytoplasmic layer around the germinal vesicle; cs, cytoplasmic 
strand; sp, sperm body. 

2. Germinal vesicle of growing micro-ovum, X 1350, camera lucida. 

C8, cytoplasmic strand; cy, cytoplasmic layer around germinal 

3. Grerminal vesicle of full-grown micro-ovum; with a sperm within, 

X 1350, camera lucida. cs, cytoplasmic strand; cy^ cytoplasmic 
layer; e, penetration path of the sperm; k, karyomerite; m, micro- 
some; sp,n, sperm nucleus. 

4. Remaining portion of germinal vesicle undergoing fragmentation, 

X 1350, camera lucida. k, karyomerite; ks, karyoplasmic 

5. Sperm body within germinal vesicle becoming vesicular preparatory 

to fragmentation, X 1350, camera lucida. 

6. Sperm body within germinal vesicle fragmenting into karyomerites, 

X 1350, camera lucida. 

7. Ovocytes differentiating from auxocytes (in banhawin), X 1350, 

camera lucida. c, centrosome; p.6i, first polar body; Ldc, intra- 
nuclear division center. 

8. Young ova in different phases of growth, X 1350, camera lucida. 

e.dc, extra-nuclear division center; Ldc, intra-nuclear division 
center; sp. sperm after penetration. 

Plate 11 

Fig. 1. Young ova, X 1350, camera lucida. sp, sperm that has penetrated. 

2. Photomicrograph of a large portion of mature ovum, X 900. Note 

sperm within germinal vesicle. 

3. Full-grown macro-auxocyte of paramamosain, in the contraction- 

figure, X 1350, camera lucida. c, centrosome with some cen- 

352 The Philippine Journal of Science *^® 

4. Diffused stage, tending toward diminution of karyoplasmic mate- 

rial preparatory to differentiation of the ovocyte, X 1350, camera 
lucida. c, centrosome. 

5. Second synizesis, or contraction-figure, showing differentiation of 

ovocyte from auxocyte body, X 1350, camera lucida. 

Plate 12 

Fig. 1. Micro-auxocyte, in contraction-figure, or synizesis, X 1350, camera 
lucida. Nuclear material undergoing rearrangement prepara- 
tory to differentiation of ovocyte. 

2. A contraction-figure in advance stage; nucleus undergoing abortive 

division, X 1350, camera lucida. c, centrosome. 

3. Micro-ovocyte that has differentiated from auxocyte body,^ X 1350, 

camera lucida. p,fu polar nucleus. 

4. Photomicrograph of portion of ovary of paramamosain, X 900. 

/, follicular wall; mi, micro-ovum; spi, sperm outside of egg; 
spat sperm within egg but still outside the germinal vesicle; sqs, 
sperm inside germinal vesicle; v, vitellogenous layer. 

5. Auxocyte of S. serrata (from an immature individual) showing 

contraction-figure, or synizesis, after growth period, X 1350, 
camera lucida. 

6. Ovocyte differentiating from auxocyte body, X 1350, camera lucida. 

pM, polar nucleus. 

7. Ovocyte wherein processes of concentration and contraction are 

apparently going on, X 1350, camera lucida. c, centrosome with 
some centroplasm; p.n, polar nucleus. 

8. Ovocyte in contraction phase, showing sloughing-off of some mate- 

rial, X 1350, camera lucida. p.tii, first polar nucleus; p.Uz, 
second polary body. 
9-10. Chromosomal constitution, X 1350, camera lucida. x, apparently 

Plate 13 

Fig. 1. Egg undergoing cleavage (S. tranquebarica) , X 900, camera 
lucida. 1, presumptive head region; 2, presumptive dorsal region; 
3, ventral region; ch, chorion membrane; 6, plasma block; p, 
blastema; p.s, perivitalline space; s.f, secondary yolk-cleavage 
furrow; v, vitelline membrane. 
2. Eggs in different stages of development, still attached to the en- 
dopodite setie, X 150, camera lucida. c, collagenous material 
attaching egg to seta; ch, chorion membrane; p.6, polar body; s, 
seta; st, stalk drawn out from layer of collagenous material 
around egg. 

Plate 14 

Fig. 1. Outline drawing of embryo, showing growth differences between 
cephalothorax and abdomen, X 900, camera lucida. Only plasma 
blocks along periphery are drawn. c,l, caudal lobe; ch, chorion 
membrane; cm, layer of collagenous material; p,l, procephalic 
lobe; st, stalk formed from collagenous material that enables 
embryo to attach to endopodite set«; th. I, thoracic lobe. 

7«*s Illustrations 353 

2. Early protozoea, X 150, camera lucida. o6, abdomen; e, eye; 

t, telson. 

3. Later protozoea stage, X 150, camera lucida. a-i, antennule; a-^, 

antenna; e, eye; m, mandible; mx-l, first maxilla; mx-2^ second 
maxilla; mp-l, first maxilliped; mp'2, second maxilliped; t, tel- 
son; 1-5, abdominal segments. 

EsfAWPAiioK: .^TfjiiioH ON 8fehm,\t«cjknksis. ] 

IPlllLH*. .JoTK. SCI.. 78, Kn. 3. 



( "•arJ 


Estampador: Studies on Spermatogenesis.] 

[Philip. Jour. Sci., 78, No. 3. 



[Philip. Jour. Sci., 78, No. 3. 






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Estampadok: Studies on Spermatogenesis.] 

[Philip. Jour, Sci., 78, No. 3. 



[Philip. Jour. Sci., 78, No. 3. 


Estampador: Studies on Spermatogenesis.] 

[Philip. Jour. Sci., 78, No. 3. 


Estampador: Studies on Spermatogenesis.] 

[Philip. Jour. Sci., 78, No. 3. 



[Philip. Jour. Scr., 78, No. 3. 







[Philip. Jour. Sci.. 78, No. 3. 


Estampador: Studies on Spermatogenesis.] 

[Philip. Jour. Sci., 78, No. 3. 




■•^Tx i-.^. ,. 


PLATE 10. 


[Philip. Jour. Sci., 78. No. 


PLATE 11. 


[pjfii.ip. Joim. Sci., 78, No. 3. 






PLATE 12. 


[Philip. Jour. Sci., 78, No. 3. 


PLATE 14. 



By Mariano P. Ramiro * 

Formerly, Fellow, Office of International Information and 

Cultural Affairs, Department of State, UJS.A. 

Abaca (Manila Hemp) waste is available in large supply in 
the Philippines and is a source of useful fibers that could be 
converted into valuable products of conunerce and industry. 
This investigation was carried out to determine the most suit- 
able conditions for reducing this material into pulp of good 
yield, strength and color. 

The term *'manila hemp" is commonly applied to the fiber 
extracted from the leaf-stalks of Mitsa textilis Nee (family 
Musacex), a plant indigenous to the Philippines. The plant 
which is locally called "abaca/' shoots from its underground 
rootstock spurious stems consisting of concentric sheating 
leafstalks. The stalk attains a height of 15 to 20 feet in 1% 
years and spreads out into large undivided leaves. Clusters of 
from 12 to 20 stalks are developed from each rhizome. In its 
native habitat, particularly the southern regions of Luzon, abaca 
grows well with the least cultivation and care. 

"Abaca" is sufficiently matured for stripping at flowering 
age. The stalk is cut down and the outer fiber-bearing layer 
of each successive leaf -stalk is divided or torn into ribbons of 
from one to two inches wide and about one-fourth of an inch 
thick. While fresh and succulent, the ribbons are scraped with 
a knife-block stripping device to render the fiber free of the 
soft cellular materials that surround the fiber strands. The 

* The author wishes to extend his appreciation to the Office of Inter- 
tional Information and Cultural Affairs of the U. S. Department of State 
for its support in carrying out this work; to the Forest Products Labora- 
tory, Madison, Wisconsin, for furnishing the facilities; to John Manning 
Paper Company, Inc., for furnishing the information on pulping Manila 
hemp; and to Messers, Mark W. Bray and John S. Martin, Forest 
Products Laboratory, for their advice and assistance in planning the 
experiments herein reported. 

9548 e 355 

356 ^^^ Philippine Journal of Science ^^^ 

extraneous material is called abaca bagasse or the plantation 
waste of the abaca industry. 


Acid sulfite pulping of abaca bagasse waste ^ results in the 
retention of most of the cellular constituents in the pulp. This 
type of pulp is suitable only for paper where strength is of 
primary consideration, but not paper for which color is essential. 

The conventional manila hemp for rope manufacture, and to 
some extent as base material for plastic articles, is usually 
clean and of high quality. It is perhaps on this account that 
the pulp yield from old manila rope by the caustic soda, soda 
ash and lime-soda ash processes is higher than the yield from 
rope factory waste ^ and low grade abaca fiber. Papers made 
from the factory waste are found also to be inferior to those 
from old rope stock. 

By digesting manila hemp with a strong solution of caustic 
soda at 320 degrees Fahrenheit Osborne 2 obtained non-hy- 
drated pulp which was converted into thin paper suitable for 
stencil base, condenser, and tissue paper. Shaw and Bicking ^ 
found that cooking wth soda ash gave satisfactory results as 
to pulp yield and quality of paper from rope stock. 

The use of abaca in the paper industry dates back to the 
American civil war^ when the production of cotton fell so 
low that the price of cotton bagging which was used exclusively 
for transporting flour and other articles prior to the war became 
prohibitive. Flour bag substitute was then an urgent problem. 
It was not very long, however, before paper of exceptional 
strength and durability was prepared from old rope stock to 
replace the cotton bag. The paper known conmiercially as 
manila paper has been so successfully manufactured from old 
rope that it has become an important material for the transpor- 
tation of food articles, cement and other commodities. The 
convertibility of manila hemp into various types of paper is 
responsible for the increasing interest and demand of the fiber 
by many industries. 


The sample of abaca waste, consisting of fifteen bales weigh- 
ing about 150 pounds each which was used for these pulping 
experiments, came from Manila and was received at the Forest 
Products Laboratory, Madison, Wisconsin, in May, 1946. A 

^*' ^ Ramiro: Suitability of Manila Hemp 357 

history of the fiber was not received. However, it appeared to 
be a fairly good rope factory waste, which fell under the T3 
quality based on the grading practice of the cordage industry. 
Besides being dusty, some of the fibers were so badly decayed 
that they crumpled on handling. Leaves, twigs, and shrub 
stalks, estimated at approximately one per cent, constituted some 
of the foreign materials mixed with the fibers. The presence of 
these undesirable materials, particularly the decayed portions 
of the hemp, contributed undoubtedly to the low pulp yield. 


Preparation of material. — ^In order to eliminate as much 
as possible the effect of the decayed fibers on pulp yields, only 
sound material was used in the pulping experiments. Twigs 
and other foreign matters were also removed but some escaped 
examination and finally emerged from the digested material 
as screenings. To overcome the difficulty of sampling and 
packing the fiber into the digester, the original fiber was cut 
into convenient sizes of from 2 to 4 inches long. 

Equipment. — Digestion experiments were carried out in 
three autoclaves each consisting of two hollow concentric cast- 
steel spheres mounted on trunnions. Each autoclave had a 
convenient opening for charging and discharging and was 
provided with a removable ground-in cover held in place by a 
suitable clamp attached to the cooking vessel. Between the 
spheres there was an annular space serving as a steam jacket 
for heating the inner sphere. One trunnion was used as an 
inlet for the admission of steam, or of cooling water to the 
jacket which may be drained through a pet cock. The other 
trunnion which projects to the center of the inner sphere 
served as a thermometer well and also supported a guard for 
the projecting steam of the thermometer. 

Each autoclave was driven by a i h. p. geared motor at 
about one revolution per minute. To the J inch steam line 
leading to the cooking vessel, about a foot from the end of the 
trunnion, a pressure gauge is installed as a means of showing 
the pressure of the steam in the steam jacket. 

Chemical — For each digestion, an equivalent amount of 
the chemical required for a cook was taken from a relatively 
strong stock solution containing 118 grams of sodium hydroxide 
per liter. 

Procedure. — ^A sample equivalent to the moisture-free 
weight of two pounds of the fiber was used in all the digestion 

358 ^^^ Philippine Journal of Science *^^ 

experiments. This quantity of material, when properly packed 
was sufficient to charge one autoclave to its full capacity of 3.7 
gallons (1/2 cu. ft.). The fiber was placed in the digester and 
the cooking liquor diluted to the proper concentration was added 
to it. For all the experimental tests, the ratio of the volume 
of the cooking solution to the moisture-free weight of the fibrous 
material was 7 to 1. The digesters were rotated throughout 
the entire cooking period. 

During the temperature rise period of one hour, steam was 
admitted to the jacket at a rate sufficient to effect a uniform 
rise of temperature. One set of pulping experiments was made 
at a maximum temperature of 140° C, another at 150° C, and 
a third at 160° C. 

In order to gain an idea of the effect on pulp yield, strength, 
color of the pulp of digestion temperature and ratio of chemical 
to raw material, a series of ten cooks was made. The liquor- 
hemp ratio and the temperature rise schedule were maintained 
constant. The length of the cooking period was also held cons- 
tant except in certain instances where it was desired to determine 
the progress of the reaction under a particular set of conditions. 
At the end of the digestion, the steam pressure in the jacket was 
relieved and the autoclave was cooled by allowing cold water 
to replace the steam in the jacket. The digested material was 
discharged in a wire-screen-bottom box where the pulp was tho- 
roughly washed. Although the digested material was consi- 
derably softened, the fiber bundles were still intact. These 
were broken up by agitating the raw cooked pulp with a suf- 
ficiently large amount of water in a tank provided with an im- 
peller stirrer. The resulting pulp was screened through a one 
foot square flat vibrating screen with slotted opening of about 
0.014 inch wide. The total yield of pulp, the screenings, and 
the screened pulp were calculated from the moist weight and the 
moisture content of the materials. 

The physical characteristics of the pulp were determined ac- 
cording to the standardized procedure and testing practice of the 
Technical Association of Pulp and Paper Industry. The average 
values obtained from the tests are a measure of the quality of 
the pulp. 


The data presented in Table 1 show the pulping conditions 
employed, pulp yields, and the physical properties of the pulps. 
In general the strength properties of the pulps were slightly 

'^ ^ Ramiro: Suitability of Manila Hemp 359 

affected by the several digestion variables including temperature 
of digestion, concentration of cooking liquor, chemical ratio, 
and time of cooking. It is apparent that in cases where a 
light-colored paper is not required, pulps from reasonably 
mild digestions are almost identical to and consequently just 
as good as the products obtained under more drastic cooking 
conditions. Incidentally, the data tend to show that pulping 
abaca waste for two hours at a maximum temperature of 150*" 
C. with from 12 to 16 per cent caustic soda (cooks 1712 and 
1713), based on the moisture-free weight of the raw material, 
give satisfactory pulps of good bursting, tearing, folding, and 
tensile strengths. The color of the unbleached pulps obtained 
under these conditions has a brightness of from 22 to 29 per cent 
as measured with the Hunter Reflectometer using a blue glass 
filter and calibrated according to TAPPI Method T-217M45. 

An increase in either the concentration of chemicals in the 
cooking liquor, the chemical ratio, the temperature, or the dura- 
tion of cooking resulted in a decrease in the permanganate 
number of the pulps. Pulps with low permanganate number 
generally have a low bleach requirement. 

In order to determine the most suitable conditions for con- 
verting the hemp into pulp with reasonably good physical prop- 
erties, batches of the raw material were digested under a number 
of cooking conditions using several chemical ratios, concentra- 
tions, temperatures and cooking times. For the first trial, cook 
1712, an initial set of cooking conditions was arbitrarily se- 
lected as a guide for the subsequent digestion experiments. 
The results show that the screenings were relatively low, an 
indication that the chemical used was almost sufficient for the 
complete reduction of the fiber to pulp. A reduction of the 
chemical ratio from 12 to 8 per cent (cook 1723) resulted in a 
decrease of from 54.7 to 48.3 per cent in the yield of screened 
pulp and an increase in screen rejects of from 1.9 to 22.4 per 
cent. Conversely, as the chemical ratio was increased from 
12 to 20 per cent at constant volume and the solution concentra- 
tion increased from 17.0 to 29.0 per cent there was a correpond- 
ing decrease in the yield of screened pulp. 

A correlation of screened yield and total yield, as affected 
by the chemical ratio and solution concentration, is indicated 
in Fig. 1. It will be noted that in the regions of low concentra- 
tion, the two curves are divergent but they converge at the 
higher concentrations. 


The Philippine Journal of Science 










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78, 3 

Ramiro: Suitability of Manila Hemp 


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362 The Philippiite Journal of Science *^^ 

The pulps from cooks 1712, 1713 and 1724 when processed in 
the test beater for 20 minutes developed a bursting strength of 
1.28, 1.46 and 1.42 points per pound per ream, respectively, 
and a tensile strengths of 6040, 6680 and 7410 lbs. per square 
inch, respectively. A tear of 2.42 grams per pound per ream 
was attained for cook 1712, 2.15 grams per pound per ream 
was obtained from 1713, while 2.85 grams per pound per ream 
was reached in 1724. The small range of variations in the 
physical properties of the pulp due to a change in chemical 
ratio and concentration of the cooking liquor tends to show 
that pulping abaca waste could be satisfactorily accomplished 
under mild conditions provided the color of the pulp is not 
considered important. Pulp yields from mild cooking conditions 
are generally higher than those obtained from a more drastic 
type of digestion. Both high pulp yields and low chemical 
requirements, associated with mild cooking conditions, are ad- 
vantageous from the point of view of conversion economy. 

Maintaining other variables constant, a difference of from 
1 to 2 hours in the cooking schedule had no distinct effect on 
the yield of screened pulp (cooks 1713, 1751 and 1752). With 
an increase of one hour in the time of digestion (cooks 1751 
and 1713), however, a decrease of approximately 1.3 per cent 
in the screenings on the basis of the moisture-free weight of the 
material was noted. The total yields from cooks 1713 and 
1751 were equal and differ from that of cook 1752 (51 per 
cent) by 0.8 per cent. The brightness of the pulp from the 
one hour digestion was slightly higher than the color of the 
pulps from cooks of longer duration. The pulp obtained from 
a two-hour digestion (cook 1713) developed, after 20 minute 
beating, a burst of 1.46 points, a tearing resistance of 2.15 
grams per pound per ream and a tensile strength of 6680 pounds 
per square inch as compared to a burst of 1.23 points, a 
tearing resistance of 2.38 grams per pound per ream and a 
tensile strength of 6800 for a one-hour digestion (cook 1751). 
On the other hand, a three-hour cook (1752) gave a burst of 
1.29 points, a tearing resistance of 2.10 grams per pound per 
ream and a tensile strength of 7580 pounds per square inch. 
From the standpoint of screened jdeld and physical properties 
of the pulps, the two-hour digestions at a maximum temperature 
of ISO^'C. with chemical ratios of 12 and 16 per cent on the 
moisture-free weight of the material were suitable for reducing 
abaca waste to pulp. 

tS, 3 

Ramiro: Suitability of Manila Hemp 


The significance of digestion temperature on pulp yields and 
on the physical properties of the pulp may be noted from the 
results obtained from cooks 1733, 1713, and 1735, which were 
digested at 140°C., 150°C. and 160°C., respectively. It was 
observed that there was a little difference in the yields of scree- 
ned pulps when the temperature was increased from 140°C. to 
160°C., although the screenings were reduced from 2.4 to 0.2 
per cent. Cook 1713 which was digested at a maximum tempe- 
rature of 150° C. gave the best yield of screended pulp and 
good physical properties which were practically indentical to 
those obtained from cook 1735 which was digested at a maximum 
temperature of 160° C. A pulp resulting from a low digestion 
temperature (cook 1733) seemed to exhibit low bursting strength 
compared to the strengths of pulps prepared at the higher 

Table 2. — Physical properties of manila paper and hand sheet. 






Experimental hand sheet (cook 1734) . 

Pts, per 

per ream 



per lb. 

per ream 


Lb, per 





Commercial manila paper * 

* Forest Products Laboratory Project 7253-J4. 

The physical properties of one of the experimental pulps 
(cook 2734) beaten for 20 minutes are compared with those 
of a commercial manila paper in Table 2. In contrast to the 
identical tensile strengths of the two samples, the folding en- 
durance of the experimental pulp, which could be considered 
satisfactory^ is one half of that of the commercial manila paper. 
On the other hand, the bursting strength of the commercial 
sample is a little lower than that of experimental test sheet. 


1. Abaca waste can be easily reduced to a suitable soda 
pulp under mild condition of digestion. The production of 
lighter colored pulps, however, requires slightly more drastic 
cooking conditions and employing higher temperatures and 
liquor concentrations. 

2. Abaca can be satisfactorily reduced to pulp by cooking 
for two hours at a maximum temperature of 150°C. with 
from 12 to 16 per cent caustic soda (NaOH) based on the 
moisture-free weight of the material. 

364 The Philippine Journal of Science ^^^ 

3. The rate of hydration on beating soda pulp from abaca 
waste is relatively fast. The pulp is sufficiently hydrated in 
20 minutes to give a Schopper-Riegler f reeness within the range 
of 550 to 700 c.c. and an average bursting strength of 1.33 points 
per pound per ream. 

4. The lightest colored pulp had a brightness of 36.4 per 
cent. The color of most of the pulps was similar to that of 
kraft pulps made from wood. 


1. Chidester, G. H. The beating of Manila hemp rope stock in the 

Rod Mill.. Forest Products Laboratory Project 253~J4. 

2. OsBORNEy F. H. Porous long fibered nonhydrated paper. Patent 

2,045,096 (November 26, 1934). 

3. Richmond, G. H. Philippine fibers and fibrous substances: their suit- 

ability for paper making Part III. Philip. Jour. Sci. 2 No, 2; 
(May, 81-107. 1907) 

4. Saleeby, M. M. The manufacture of paper from abaca. Philip. Agri. 

Rev. (1913), 23-27. 

5. Shaw, M. G. and G. W. Bicking. Manila rope waste for paper 

manufacture. Paper Trade Journal 84, No. 18 j (May 5, 68-70 


Books reviewed here vi^ere received from time to time by 
the Philippine Journal of Science and acknowledged in this 


Sergey's Manual of Determinative Bacteriology. By R, S. Breed, E. G. 
D. Murray and A. Parker Hitchens. Sixth edition. Baltimore, The 
Williams & Wilkins Company, 1948, 1529 pp. Price $15.00. 

The sixth edition of Bergey's Manual of Determinative Bac- 
teriology is to the present the most complete study of taxonomy 
and nomenclature of bacteria. The work is accomplished by the 
permanent nomenclature committee created by the International 
Society of Microbiologists. 

The nomenclature used in the latest edition diif ers from that in 
the previous edition. In the present edition the nomenclature 
is based upon the similar antigenic structures which identify 
the serotypes rather than the species. The inclusion of the SaU 
monella typhosa in the Genus Salmonella, called formerly Ba- 
cillus typhosus and Eberthela typhosa, is an example. 

Besides the committee, sixty contributors assisted in the 
preparation of this edition. Since this book is the product of 
extensive study of renowned bacteriologists and specialists it 

^*' ' Ramiro: Suitability of Manila Hemp 365 

is therefore highly recommended as a valuable reference 
material for all scientific institutions and researchers. The 
duplication of names of certain species of bacteria has been 
rather confusing to any one who is unaware of the existence of 
certain synonyms. The publication of this manual has solved 
that confusion. Furthermore, this book also gives the general 
rules of nomenclature and principles of taxonomy so that a 
student of bacteriology may readily recognize the terms used 
regularly in classification. Following the rules outlined in this 
book it is easy for the students to identify a certain microor- 
ganism which they may encounter in their research work. 

The book is really an invaluable help. People interested in 
bacteriology should consider its acquisition as a real and per- 
manent investment. — ^M. B. 

Biochemical Preparations. By Advisory Board, W. Mansfield Clark and 
others; Editorial Board, Herbert E. Carter, Ed.-in-Chief. New York, 
John Wiley & Sons, Inc., ^949. v.l. Price $2.50 

This book is the first volume of what is to be hoped a rich 
and comprehensive contribution to detailed methods of the 
preparations of various substances of biochemical origin. 

It follows the same trend as the series of Organic Synthesis 
and Inorganic Synthesis series — ^but as the title indicates the 
book gives detailed methods as to the separation and purifica- 
tion of certain biochemical products from natural sources, as 
well as synthesis of some from organic compounds. 

Methods are claimed to be precise as they have been counter- 
checked.— F. M. Y. 

Experimental Physics for Colleges. By Walter A. Schneider and Lloyd 
B. Ham. Revised edition. New York, The Macmillan company, 1949. 
442 pp., illus. Price, $3.80. 

The book is a revised edition of the text by the same authors 
on experimental physics for elementary students. 

The new edition is, as the title indicates, intended for use in 
colleges . It is composed of 7 parts : Part 1, Units and Measure- 
ments; Part 2, Mechanics; Part 3, Sound; Part 4, Heat; Part 5, 
Electricity; Part 6, Light; Part 7, Notes and Tables. There 
are 34 chapters, 56 experiments and 12 standard tables. An 
index is included. 

The authors succeeded in this new revised edition, in present- 
ing the subject of experimental physics for colleges, in a clear 
and concise manner, particularly to those who want to pursue 
advanced studies in physics. The mathematical treatment is in 

366 The Philippine JoumaX of Science *^^ 

accord with those covered in the first year of college. Figures 
are included to help the reader in understanding the different 
principles involved in the experiments. Like any other book 
of this kind, numerous problems are given to increase its 
usefulness to both the instructors and students. The chapter 
on Notes and Useful Instruments and Procedures is a good 
addition to the book. As a whole, the book is recommendable 
as a reference or as a text to those interested in somewhat ad- 
vanced treatment of experimental physics, especially those in 
colleges. — G. O. 0. 

The Philippine 
JouBNAL OF Science 

Vol. 78 DECEMBER, 1949 No. 4 


On October 26, 1949, Dr. Marcos A. Tubangui, our foremost 
Filipino parasitologist of international renown, passed to the 
Great Beyond after a short illness. His passing is a great loss, 
not only to the Philippines, but also to the whole science of 
parasitology; it left a void which will be difficult to fill for many 
years to come. 

Dr. Marcos A. Tubangui was born in Porak, Pampanga, on 
April 25, 1893. He studied in the public schools and graduated 
from the College of Veterinary Science, University of the 
Philippines, in 1918. After graduation, he served as veterina- 
rian in the Bureau of Agriculture for a year, and then was sent 
abroad as fellow of the University of the Philippines. 

While in the United States, he attended the Marine Biological 
Laboratory at Woods Hole, Massachussetts ; the University of 
Illinois; and worked at the Bureau of Animal Industry, U. S. 
Department of Agriculture. In 1921, he received his Master 
of Science degree at Cornell University. 

Upon his return to the Philippines in that year, he was made 
instructor and, later, assistant professor of veterinary parasi- 
tology, University of the Philippines. In 1931, he transferred 
to the Bureau of Science and spent the best years of his life 
as Chief of the Division of Biological Research. In 1948 he 
was appointed Professor and Head of the Department of Para« 
sitology, Institute of Hygiene, University of the Philippines, a 
position which he held until his death. 

* By Dr. T. P. Pesigan, senior parasitologist and chief, Schistosomiasis 
Control Program, Department of Health, Republic of the Philippines. 

10832 S67 

368 The Philippine Journal of Science 


Dr. Tubangui was an active member of many learned societies 
and professional organizations. He was a member of Sigma Xi, 
the American Association for the Advancement of Science, the 
American Society of Parasitologists, the Society for the Ad- 
vancement of Research, Los Bafios College, of which he was 
once Secretary, the Philippine Scientific Society, the Philippine 
Veterinary Medical Association, the Philippine Medical As- 
sociation (associate member), the National Research Council 
of the Philippines (Charter Member) , and the Philippine Society 
of Parasitology, of which he was President for many years until 
his death. 

He contributed a great deal to the science of parasitology, 
particularly taxonomy of many helminths. He wrote and pub- 
lished many scientific papers on both medical and veterinary 
parasitology. Most outstanding of his contributions to medical 
parasitology are the complete life cycle of Euparyphium ilocanum 
which he worked out, and his discovery of the molluscan interme- 
diary host of the Schistosoma japonicum in the Philippines. 
With these investigations, he paved the way towards the control 
of these parasitic diseases in this country. He also left several 
unfinished works, especially on paragonimiasis and filariasis, 
which even a few days before his death, he repeatedly told his 
wife he would wish to continue, still hoping he would get well. 

The career of Dr. Tubangui is a great inspiration to the 
young researchers of the Philippines; for as a leader in his 
field, he was always helpful and kind to his younger associates. 
They, in turn, loved and respected him. Above all, he was 
always a silent and devoted worker. He sustained his enthu- 
siasm for research until his death, despite diflJculties. His deep 
understanding of human nature and his undying interest in 
his work always kept him away from controversies. He died 
a true scientist in the strict sense of the word. 


Tubangui, M. A. Two new intestinal trematodes from the dog in China. 

Proc. U. S. Nat. Mus., Art 2415, 60 (1922) 1-12. 
Tubangui, M. A., G. San Agustin and F. M. Foronda. Parasitological 

studies by the use of collodion sacs implanted intraperitoneally. 

I. Notes on the life history of Ascaris lumbricoides, Philip. Agric. 11 

(1922) 153-158. 
Tubangui, M. A. and B. Schwartz. Uncommon intestinal parasites of 

man in the Philippine Islands. Reports of new cases. Philip. Jour. 

Sci. 20 (1922) 611-618. 
Tubangui, M. A. The prevalence of hookworm and other intestinal 

nematodes in adult Filipinos. Jour. Parasitol. 9 (1922) 83-92. 

^'^' * Pesigan: Marcos A. Tuhangi 369 

TuBANGUi, M. A. Parasites of lower animals dangerous to man in the 

Philippine Islands. Philip. Agric. 11 (1922) 243-250. 
TUBANGUI, M. A. Two larval parasites from the Philippine palm civet 

{Paradoxurus philippinensis) . Philip. Jour. Sci. 24 (1924) 749-755. 
TUBANGXJI, M. A. Metazoan parasites of Philippine domesticated animals. 

PhiUp. Jour. Sci. 28 (1925) 11-37. 
TuBANGUi, M. A. and S. A. Francisco. Report on the hookworm campaign 

carried among the students of the Los Banos Colleges during the 

collegiate year 1924-1925; Ann. Rep. Dean Coll. Agric, Univ. 

Philippines for the Year 1924-1925, 144-148. 
TuBANGUI, M. A. Worm parasites of Philippine chickens. Philip. Agric. 

Rev. 19 (1926) 1-43. 

TUBANGUI, M. A. Larval trematodes from Philippine snails. Philip, Jour. 
Sci. 36 (1928) 37-54. 

TUBANGXH, M. A. Trematode parasites of Philippine vertebrates, I. Philip. 
Jour. Sci. 36 (1928) 351-371. 

TuBANGUl, M. A. Paradistomum gregarinum, a new name for the trema- 
tode Paradistomum magnum. Philip. Jour. Sci. 38 (1929) 443. 
TuBANGUi, M. A. and E. C. Farinas. Two tapeworm parasites from the 

carabao, with special reference to a new species of Avitelina, Philip. 

Jour. Agric. 1 (1930) 421-429. 
TUBANGUI, M. A, and S. A. Francisco. The presence in human stools of 

the eggs of a trematode parasitic in jfish. Jour. Philip. Is. Med. 

Assoc. 10 (1930) 31-33. 
TUBANGUi, M. A. Trematode parasites of Philippine vertebrates. II: Two 

echinostome flukes from rats. Philip. Jour. Sci. 44 (1931) 273-283. 
TUBANGUI, M, A. Eimeria hukidnonensis, a new coccidium from cattle, 

and other coccidial parasites of domesticated animals. Philip. Jour. 

Sci. 44 (1931) 253-271. 
TuBANGUl, M. A. Worm parasites of the brown rat (Mils norvegicua) in 

the Philippine Islands, with special reference to those forms that 

may be transmitted to human beings. Philip. Jour. Sci. 46 (1931) 

TuBANGUl, M. A. Trematode parasites of Philippine vertebrates, III: 

Flukes from fish and reptiles. Philip. Jour. Sci. 44 (1931) 417-423. 
TUBANGUI, M. A. Trematode parasites of Philippine vertebrates, IV: 

Ectoparasitic flukes from marine fishes, Philip. Jour. Sci. 45 (1931) 

TuBANGUl, M. A. and R. Villamil. Nematodes in the collection of the 

Philippine Bureau of Science, I: Oxyuroidea. Philip. Jour. Sci. 51 

(1931) 607-615. 
TUBANGUI, M. A. and L. M. YuTUC. The resistance and the blood sugar 

of animals infected with Trypanosoma evansL Philip. Jour. Sci. 45 

(1931) 93-107. 
TUBANGUI, M. A. Themolluscan intermediate host in the Philippines of 

the Oriental blood fluke, Schistosoma japonicum Katsurada. Philip. 

Jour. Sci. 49 (1932) 295-304. 

370 The Philippine Journal of Science i»49 

TUBANGUI, M. A. Trematode parasites of Philippine vertebrates, V : Flukes 
from birds. Philip. Jour. Sci. 47 (1932) 369-404. 

TUBANGUI, M. A. Observations on the life histories of Euparyphium 
fnurinum Tubangui, 1931, and Echinostoma revolutum (Froelich, 1802) 
(Trematoda). Philip. Jour. Sci. 47 (1932) 497-413. 

Tubangui, M. A. Observations on the possible transmission of surra by 
the land leech, Haemadipsa zeylanica, Philip. Jour. Sci. 4S (1932) 

Tubangui, M. A. Notes on Acanthocephala in the Philippines, Philip. 
Jour. Sci. 50 (1933) 315-128. 

Tubangui, M. A. Trematode parasites of Philippine vertebrates, VI: 
Descriptions of new species and classification. Philip. Jour. Sci. 
52 (1933) 167-197. 

Tubangui, M. A. and A. M. Pasco. The life history of the human 
intestinal fluke, Euparyphium ilocanum (Garrison, 1908) Philip. Jour. 
Sci. 51 (1933) 581-606. 

Tubangui, M. A. Nematodes in the collection of the Philippine Bureau 
of Science, II: Filarioidea. Philip. Jour. Sci. 55 (1934) 115-123. 

Tubangui, M. A., M. Basaca and A. M. Pascx). Hexylresorcinol as an 
anthelmintic: its efficiency against the intestinal parasites of man, 
Philip. Jour. Sci. 54 (1934) 473-481. 

Tubangui, M. A., and L, M, Yutuc. Human infestations with Ascaris 
and Trichuris in different parts of the Philippine Islands. Philip. 
Jour. Sci, 55 (1934) 91-113. 

Tubangui, M. A. Additional notes on Philippine Acanthocephala. Philip. 
Jour. Sci. 56 (1935) 13-19. 

Tubangui, M. A., M. Basaca, A. M. Pasco and F. del Eosario. Observa- 
tions on the geographical distribution of hookworm parasites and 
hookworm disease in the Philippines. Philip. Jour. Sci. 58 (1935) 

Tubangui, M. A. and V. A. Masilungan. Trematode parasites of Philip- 
pine vertebrates, VII: Additional records of new species. Philip. 
Jour. Sci. 58 (1935) 435-445. 

Tubangui, M. A. and V. A. Masilungan. Trematode parasites of Philip- 
pine vertebrates, VIII: Flukes from a cobra and a crocodile. Philip. 
Jour. Sci. 60 (1936) 255-265. 

Tubangui, M. A. and V. A. Masilungan. Studies on the cercariacidal 
property of the sera of vertebrate animals. Philip. Jour. Sci. 60 
(1936) 393-398. 

Tubangui, M. A. and V. A. Masilungan. Notes on Philippine linguatulids 
(Arthropoda: Pentastomida). Philip. Jour. Sci. 60 (1936) 399-405. 

Tubangui, M. A. and V. A. Masilungan. Oochoristica excelsa, a new 
reptilian cestode. Philip. Jour. Sci. 61 (1936) 75-79. 

Tubangui, M. A. and V, A. Masilungan. Diplosentis amphacanthi gen. 
et sp. nov., an Acanthocephala parasitic in a marine fish. Philip. 
Jour, Sci. 62 (1937) 183-189. 

Tubangui, M. A. and V. A. Masilungan. Tapeworm parasites of Philip- 
pine birds. Philip. Jour. Sci. 62 (1937) 409-438. 


Pesigan: Marcos A. Tuhwngi 37X 

TuRANGUi, M. A. and V. A. Masilungan. Nematodes in the collection 
of the Philippine Bureau of Science, III: Philip. Jour. Sci. 64 (1937) 

TuBANGUi, M. A. and V. A. Masilungan. Nephridiorhynchus palawa- 
nensis sp. nov., an Acanthocephalan parasite of Manis javanica 
Desmarest. Philip. Jour. Sci. 66 (1938) 1-5. 

TuBANGUi, M. A. and V. A. Masilunigan. Trematode parasites of Phil- 
ippine vertebrates, IX: Flukes from the domestic fowl and other birds. 
Philip. Jour. Sci. 75 (1941) 131-141. 

TuBANGUi, M. A. and V. A. Masilungan. Some trematode parasites of 
fishes in the collection of the University of the Philippines. Philip. 
Jour. Sci. 76 (1944) 213-221. 

TuBANGUi, M. A. and V. A. Masilungan. On two Acanthocephala from 

the Philippines. Jour. Parasitol. 32 (1946) 154-155. 
Tubangui, M. a. Pseudophyllidean cestodes occurring in the Philippines. 

Livro Jubilar Prof. Travassos, Rio de Janairo, Brazil. (1938) 489-494. 
Tubangui, M. A. and E. Y. Garcia. Clinostomum ahdoni sp. nov., a 

trematode parasite of the cat in the Philippines. Philip. Jour. Sci. 70 

(1939) 397-401. 

Tubangui, M. A. and C. M. Africa. The systematic position of some 
trematodes reported from the Philippines. Philip. Jour. Sci. 67 (1938) 
117-127. Also in Volumn Jubilare Prof. Sadao Yoshida, Osaka, 
2 (1939) 145-153. 

Tubangui, M. A. and A. M. Pasco. Studies on the geographical distribu- 
tion, incidence, and control of Schistosomiasis japonica in the Philip- 
pines. Philip. Jour. Sci. 74 (1941) 301-329. 

Tubangui, M. A. and P. J. Aguila. The treatment of Schistosomiasis 
japonica with fuadin. Philip. Jour. Sci. 75 (1941) 69-73. 

Tubangui, M. A. Preliminary report on the specific identity and life 
history of the lung fluke of mammals (Paragonimus) in the Philip- 
pines, (Abstract). Abstracts of Scientific Papers before the Con- 
ference on Medical Sciences in Commemoration of the Establishment 
of the Republic of the Philippines, December 20, 21, and 22, Manila 
(1943) 36-37. 

Tubangui, M. A. Preliminary notes on the crustacean vector of the 

mammalian lung fluke (Paragonimus) in the Philippines. Jour. 

Parasitol. 32 (1946) 152-153. 
Tubangui, M. A. Plagiorchoides potamonides (Plagiorchiidae) , a new 

trematode found in experimental rats. Jour. Parasitol. 32 (1946) 


Tubangui, M. A. A summary of the parasitic worms reported from the 
Philippines. Philip. Jour. Sci. 76 (1947) 225-304. 

Tubangui, M. A. and M. Basaca. Notes on the anthelmintic properties 
of the latex of papaya. (Carica papaya Linn.), and *'Isis" (Ficus 
ulmifolia Lam.). Philip. Jour. Sci. 77 (1947) 19-24. 

Tubangui, M. A. Schistosomiasis japonica and other helminthic diseases. 
Proceedings Fourth International Congresses on Tropical Medicine and 
Malaria, Department of State, Washington D. C, 2 (1948) 1034-1039. 

372 The Philippine Journal of Science ^^^^ 

TtTBANGUi, M. A. and B. D. Cabrera. Studies on Filariasis in the Philip- 
pine Islands. Results of a Survey in the Province of Sorsogon and 
in the New Bilibid Prison at Muntinlupa, Eizal. Acta Medica Phil- 
ippina 5 (1948) 50-56. 

TUBANGUI, M. A. and B. D. Cabrera. Studies on Filariasis in the Philip- 
pine Islands. Results of a Survey in the Province of Sorsogon and 
in the New Bilibid Prison at Muntinlupa, Rizal. II: Treatment of 
Bancroftian Filariasis with Hetrazan. Acta Medica Philippina 6 
(1949) 1-7. 

I*l'.<5if;..i\ : M \uros A. TfiiANuri. f 

I l'}ii!JP. Juriu Sri., Vhi. 78. Mn. f. 




Of the Division of Laboratories, Department of Health 
With the Technical Assistance of 

RosALiNA L, Dee** 
Of the Division of Laboratories, Department of Health 

The study of thiamine content of breast milk is important in 
this country since it is the only natural source of thiamine 
available to breast fed babies. Moreover, it has been found 
in the past that the death rate among the breast fed infants 
was higher than that of artificially fed babies. This was due 
to the fact that infantile beriberi was mostly found among 
the breast fed infants. 

Vedder and Clark(i) have found that 5 cc of fresh cow's 
milk did not prevent polyneuritis in birds on a polished rice 
diet. Gibson(2) in studying the protective power of normal 
breast milk of Filipinos against polyneuritis gallinarum has 
found that it contained not more than one-fourth the amount 
of vitamin of cow's milk. Again Gibson and Concepcion(3) in 
studying the influence of fresh and autoclaved milk on the 
development of neuritis in animals have demonstrated that 20 cc 
of fresh human milk of Filipino mothers did not prevent birds 
on a polished rice diet from developing polyneuritis. The 
observation of Clements (4) that breast fed infants frequently 
exhibit partial thiamine deficiency is of interest in this con- 
nection. He also noted that partial thiamine deficiency was fre- 
quently found among the breast fed infants in Australia and 
attributed it to suboptimal quantities of thiamine in the milk, 
resulting from insufficient intake. On the other hand, Sunda- 
rajan(5) found that the average thiamine content of milk 
among women with beriberi in India was not less than the 
values found among normal women. In view of this contradic- 
tory findings the determination of thiamine content of normal 
as well as beriberi milk is of special interest in this country in 

1 Aided by special grant from the Institute of Nutrition. 

* Chief Biochemist, XJ.S.P.H.S. Rehabilitation Program and the Depart- 
ment of Health. 

^Research Assistant under special grant in aid from the Institute of 


374 The Philippine Journal of Science ^»*^ 

order to establish the relationship, if there is any, between low 
thiamine and infantile beriberi. 


The present study is a report on the thiamine determination 
made on 140 samples of mature milk taken from both breasts 
of healthy mothers eating self-chosen diets and 21 samples of 
beriberic milk from others whose babies showed characteristic 
symptoms of infantile beriberi. The samples were taken be- 
tween 9 and 10 a.m. and in different stages of lactation. The 
total thiamine was determined by the method described by 
Kendall(6) with a slight modification and can be described as 
follows : 

Five ml. of milk is placed in each one of the four reaction ves- 
sels and then 0.25 ml. of 10 per cent freshly prepared takadias- 
taseis added. Then 0.25 ml. of 10 per cent acetic acid is added. 
Incubate for 3 hours at 37°C. At the end of the period remove 
the reaction vessels and add 1 ml. of thiamine standard solution 
to the reaction vessel No. 3. Then add 3 ml. of the oxidizing 
reagent (0.1 ml. of 1 per cent K3Fe(Cn)6 in 3 ml. of 15 per cent 
NaOH) to each of the vessels Nos. 1, 2 and 3. To vessel No. 4 
add 3 ml. of 15 per cent NaOH. Shake the vessels from end 
to end gently for 45 seconds then add to each vessel 13 ml. of 
redistilled isobutanol. Shake again gently for exactly 1| min- 
utes. Centrifuge for 5 minutes, draw off the aqueous layer. 
Pour the isobutyl layer into the cuvettes and read after ad- 
justing the reading of the fluorophotometer with the quinine 

The calculation is made as follows: The average reading of 
reaction vessels Nos. 1 and 2 less the reading of the blank 
(R.V.4) divided by the reading of the reaction vessel No. 3 minus 
the average readings of reaction vessels Nos. 1 and 2 times 20 
equals micrograms of thiamine per 100 cc milk. 


The results of the present investigation are summarized in 
Tables 1, 2, 3 and 4. 

As shown in Table 1 the mean of 140 samples of normal milk 
is 7.691 mcg.± 0.322 with a standard deviation of 3.820. On 
the other hand, the mean of 21 cases of beriberic milk as shown 
in Table 2 is 4.97 ±: 0.677 with a standard deviation of 3,0277. 


Concepcion: Human Milk Studies 375 

Table 1,— Total thiamine content of normal milk. 

Age o! milk 

No. of 



S.E. M, 

1 month.. «.. - -- 

















2 months 


3 months . , , ^ - ... .. ^ , . . ,. ., „ , , .. 


4 mont hs 


5 months. - --- . 


6 months. - 


7 months .. 


8 months _. 


9 months -. 


lO months 

11 months . ^ ..,....- ,^ ... . ... ... 



12 months . 






Table 2. — Total thiamine content of beriberic milk. 

Age of milk 

No. of 



S.E. M. 

1 month- „_ _ - - - 









2 months 


3 months. - - - 

3 35 

4 mon<"hfl - - 


5 months 






Table 3. — Total thiamine content of normal milk during the first five 


Age of milk 

No. of 




1 month- - -» . - - 










2 months-- 


8 months 


4 months . 


5 months 






Table 4. — Thiamine content of human milk as reported by various authors. 





Period of 



Thiochrome .. 
Thiochrome — 
Thiochrome — 
Thiochrome .. 







2-10 da. 
3-38 wk. 
2-10 mo. 
1-12 mo. 

100 ee 

Slater and Rial 


Roderucketal -^_ 




376 The Philippine Journal of Science ^^^^ 

Since the beriberi cases were only found between the ages one 
and five months, we cannot very well compare them with the 
normal because their ages range from one to twelve months. 
The logical comparison is to compare the beriberi cases with 
the normals from one to five months also. This is summarized 
in Table 3. The difference in the mean between the normal 
and beriberic milk, as shown in Tables 2 and 3, is 2.77. This 
difference when studied statistically was found to be significant. 
The value of t being 3.48 which is significant even at 1 per cent 


According to Slater and RialC?) the average total thiamine 
content of milk from 3 to 38 weeks is 12.7 meg. per 100 cc. as 
shown in Table 4. Roderuck et al.W found an average of 
14.8 meg. per 100 cc. from 2 to 10 months. Knott's group(9) 
found an average of 15.1 meg. per 100 ml. for mature milk. 
All these findings were higher than the result of the present 
investigation. This can only be explained by the difference 
in the thiamine content of the diet. This explanation is sup- 
ported by the high incidence of beriberi in this country. Ac- 
cording to Roderuck et ah diet showed a clean cut relationship 
to the thiamine content in human milk. 

The average concentration of thiamine in milk remained 
relatively constant after the fourth week of lactation according 
to Roderuck et al. Knott reported that the thiamine in milk 
reached the level characteristic of the subject in three weeks. 
On the other hand, Slater and Rial found a gradual increase 
in the thiamine content until the twentieth week. Our results 
cannot corroborate or disprove those findings because our 
samples were not 24-hour samples. 

Knott(9) reported some determination of thiamine content 
of breast milk from 17 women where figures range from 3-18 
meg. per 100 cc. with an average of 9 meg. She attributed 
this low thiamine content of human milk to the low content 
of thiamine in the diet since food is an important factor 
influencing the thiamine content of breast milk. The question of 
whether breast milk is adequate in meeting the thiamine re- 
quirements of infants has been answered by Knott et al when 
they stated that the minimum thiamine requirement of young 
infants (1 to 6 months of age) is approximately 200 meg. daily. 
This amount can just be met by a normal healthy infant if his 
mother's milk contains 20 or more meg. of thiamine per 100 cc. 
They suggested that 40 meg. per kilogram body weight may be 
necessary to supply the needs of the infant. This finding was 
corroborated by Holt et al!,(lO). 

'^'^''* Concepcion: Human Milk Stvdies 377 


1. One hundred forty samples of mature human milk from 
normal women taking ordinary diet have been assayed for 
total thiamine and a value of 7.691 meg. ±: 0.322 per 100 ml. 
with a standard deviation of ±: 3.820 was found. 

2. Twenty one samples of mature human milk from mothers 
whose babies showed clinical symptoms of infantile beriberi 
have been assayed for total thiamine and a mean value of 4.97 
ip 0.677 per 100 ml. milk with a standard deviation of z^ 3.0277 
was found. The difference in the mean between normal and 
beriberic milk was found to be statistically significant. The 
value of t being 3.48 which is significant even at 1 per cent 

3. The determination of thiamine content of milk may be 
of some help in the diagnosis of beriberi when the clinical 
symptoms of the disease are not quite conclusive. 

4. Because of the limited number of samples of beriberi 
milk examined, it is rather risky to give a definite conclusion 
whether the mean figures given here are of some practical value 
in the diagnosis of beriberi or not even though the difference 
between the two means is statistically significant. 


1. Vedder, E. B. and E. Clark. Philip. Jour. Sci. 7 B (1912) 423-461. 

2. Gibson, R. B. Ibid 8 B (1913) 469-473. 

3. Gibson, R. B. and I. Concepcion. Ibid. 11 B (1916) 119-133. 

4. Clements:, M. W. Med. Jour. Australia 1 (1942) 12. 

5. SUNDARAJAN, A. R. Ind. Jour. Med. Res. 29 (1941) 567-573. 

6. Kendall, N. Jour, of Pediatrics 20 (1942) 65. 

7. Slater, E. C. and E. J, Rial. Med. Jour. Australia 1 (1942) 3. 

8. RODERUCK, C. E. ET AL. Amer. Jour. Dis. Child. 70 (1945) 162. 

9. Knott, E. M. Amer. Jour. Pub. Health 32 (1942) 1013-1017. 
10. Holt, Jr., E. et al. Jour, of Nutrition 37 (1949) 53-66. 


The authors wish to express their acknowledgement to the 
Director and the Board of Directors of the Institute of Nutrition 
for the grant in aid given to us to undertake this work. We 
wish also to express our appreciation to Dr. A, Tupas, Chief, 
Department of Pediatrics, Philippine General Hospital, for sup- 
plying us with samples of beriberi milk; Dr. F. Horilleno and Dr. 
W. de Leon, Director of Laboratories, Department of Health, for 
their sympathetic cooperation and advice and to Brig. General H. 
F. Smith of the U. S. Public Health Service for the various 
help given to us during the performance of this work. 


Of the College of Veterinary Medicine, University of the Philippines 


Of the known vectors of surra, the horseflies are considered 
the most dangerous. It has been estimated that there are about 
2,000 known species of hoi*seflies in the world. Patton and 
Cragg (1913) recorded 106 Oriental species belonging to the 
genus Tabanus, two of which are known to exist in the Islands 
{Tdbanus vanderwulpi and Tahanus striatus), Schuurman 
Stockhoven, Jr. (1926) recorded 239 species in the Dutch East 
Indian Archipelago and neighboring countries in the Far East, 
of which four occur in the Philippines, namely, Tabanus striatm, 
Tabamts reducens, Tabanus flexilis and Tabanus factiosus. 
Perhaps the most comprehensive systematic study of the genus 
Tabanus in the Islands is that of Kroeber (1924) who listed 
13 species, among which are the following: Tabanus ixion, 
Tabanus univentris, Tabanus flaviventris, Tabanus rubidus, 
Tabanus fumifer, Tabanus malayensis, Tabanus philippinensis, 
Tabanus malanopygatus and Tabanus graiseoscutellatus. All 
told, fourteen species of Tabanus have been described from the 
Philippines, of which T. striatus and T. reducens appear to be 
the most common (PL 1, figs. 1 and 2). 


The study on the occurrence of horseflies was conducted on 
the grounds of the Bureau of Animal Industry, and also on 
a lot formerly occupied by the College of Veterinary Medicine 
in Pandacan, Manila. The marshy land and the zacate field 
in the immediate vicinity were also included in the survey. 
Most of the flies collected were caught from the telegraph and 
electric posts on the sides of the abandoned railway tract 
crossing the area. A random examination of the soil in this 
area revealed the presence of larvae of horseflies, suggesting 
that it is a breeding place of tabanid flies. Previous observa- 
tions have shown that the flies usually rest on the posts during 


380 The Philippine Journal of Science ^®*^ 

the middle of the day. For this reason, the collection of flies 
was made between 12:00 and 1:00 o'clock in the afternoon. 
The work was started during the Japanese occupation but since 
some of the data pertaining to that period were lost, only 
those collected after liberation are included in this paper. 

During a period of three years and two months (November, 
1945 to December, 1948), 854 tabanids were caught, of which 
480 were Tahanus striatus and 374 Tabamts reducens, the 
former exceeding the latter by 106 flies. Of the 480 Tahanus 
striatus 57 were males, the male — ^female ratio being thus 
1:7.42. In the case of Tahanus reducens, 14 were males and 
360 females, the male — ^female ratio being 1 :25.71. It will be 
noted that there were more than 4 times more males of 
Tahanus striatum than Tahanus reducens and this may account 
partly for the greater number of Tahanus striatum collected 
during the period under consideration. 

While the female flies were seen throughout the year, the 
males were absent during certain months. In the case of 
Tabanics striatum, males were not found during June and July; 
the maximum catch of 15 male flies was made during the month 
of February and the minimum catch of one fly, during May* In 
the case of Tahanus reducens, three males were collected during 
each of the months of January, October, November, two in De- 
cember and one each in February, March and May. None were 
found during April, June, July, August and September, 


Of the factors which may be responsible for the monthly 
fluctuation in the prevalence of tabanid flies, only the rainfall 
seems to have an appreciable influence. A summary of the fly 
collections during the three year period and the corresponding 
rainfall data are presented in Figure 1 and Table 1, Although 
tabanid flies were always found during the three-year period, 
it is shown they were more abundant during certain months. 
A subpeak representing 37 flies was observed in February and 
this was followed by a gradual decline up to June, when the 
smallest collection of 4 flies was recorded. Thereafter, the 
monthly average catch increased up to October when the maxi- 
mum collection of 72.6 flies was made. After this month, 
the fly-densities decreased up to the end of the year. In general, 
the data agree with those of Mitzmain (1913) who found Ta- 
hanus striatus to be prevalent from October to March in the 


Yutuc: The Prevalence of Tabanid Flies 


vicinity of Manila. On the other hand, Kelser (1927) observed 
that during 1926 Tabanus striatus commenced to make its ap- 
pearance in and around Manila in appreciable number during the 
later part of July and that by the middle of August great num- 
bers of flies were in evidence. Manreza and Mondoiiedo (1935), 
working in Los Baiios, Laguna, also recorded a monthly fluc- 

A/i/mJber of Tabantds 
t \ % % % 1 % 

tA N 

< 18. i 

1 " 

I \ 


1 » § 1 

\ \ \ \ 










































Fia,l, f/f//t/e/fce of ramfcf// o/r/^e numertca/ -f/uch/afion of 
fabamcf f//ej. 

tuation in the density of Tabanus striatus. They made their 
maximum collection in May, the minimum in October. It thus 
appears that the monthly prevalence of horseflies varies consider- 
ably in different localities and possibly during different years. 
Correlating the monthly collections with the seasonal distribution 
of the annual rainfall, as shown in Figure 1, it is seen that 


The Philippine Journal of Science 


the peak of fly incidence in October occurs two months after 
the heaviest rainfall in August. In British East Africa 
Leese (1914) observed that species of Tabanus are abundant 
during and after the rainy season, the numbers falling off greatly 
as the dry season advances. 

Table 1. — Data on rainfall and its relation to tabanid incidence 






Average for 

three-year period 




m mm. 

12 3 



2 6 


10 7 


63 6 


127 1 






October — 
December - 

Average for 
three-year period 


29 3 

in mm. 



Since no records are available that Tabanus reducens has 
been found guilty as a vector of Trypanosoma evansi, it seemed 
desirable to test this horsefly in the laboratory. For compar- 
ative purpose, Tabanus striatum was also used in the transmission 
experiments. Guinea pigs and white rats kept in a fly-screened 
enclosure were utilized. The horseflies used were selected from 
those collected during the survey. It may be noted here that 
only a small percentage of the flies could be induced to bite 
under experimental conditions and could not be kept alive in 
the laboratory for long periods of time. In one instance, how- 
ever, a Tabanus reducens was maintained in captivity for a 
period of one month. 

The feeding of flies was carried on in small wire baskets 
applied on surra-infected guinea pigs where the hair had been 
previously clipped close to the skin. As a rule, the flies were 
allowed to feed for from 30 seconds to one minute, after which 
they were transferred to a clean white rat where the hair had 
also been previously shaved. The intervals between feedings 
ranged from 20 seconds to one and one-half minutes. In some 
of the flies four to five interrupted feedings could be carried 
on before they refused to bite any further. 

For convenience, the results are summarized in Table 2. 
Of four experiments with Tabanus reducens, three were positive. 
With Tabanus striatus, only one positive result was recorded 


Yutue: The Prevalence of Tabanid Flies 


out of the four trials and it was due to the use of eight flies in 32 
interrupted feedings. Mitzmain ( 1914) had practically the same 
experience with T. striattcs, for in his experiments he obtained 
only three positive results out of 16 trials. It thus appears 
that Tahanm redueens is a more efficient transmitting agent 
of the surra parasite than Tabanus striatus. Since the method 
of transmission is purely mechanical, it is most probably 
the larger size of the fly and its mouth parts which make it 
a more dangerous vector. Measurements made from a number 
of specimens show that Tabanus redueens is approximately one- 
third longer and broader than Tabanus striatus and its mouth 
parts are correspondingly bigger. 

Table 2. — Data on the mechanical transmission of surra by T. redueens 

and T. striatus 


Density of trypanosome 

in peripheral blood of 

surra infected guinea 

pig per cu. mm. 

Species and number of fiy used 

No. of 















T. redueens 2 

do 1 

do 2 

do 2 

T. striatus . Bf 

do 2 

do 8 

do _ 2 







With T. redueens there are at least 22 species of tabanid 
flies which have been proven experimentally to transmit Trypa- 
nosoma evansi. The others, according to the investigations of 
Sergent, Ed. and Et. (1905 b and 1906 a), Eraser and Sy- 
monds (1908), Mitzmain (1912), Kelser (1927), Cross and Patel 
(1922), and Nieschulz (1926, 1927, 1929, and 1941), are: 
T. albimidus, T. albitriangularis, T. bilateralis, T. brunipes, T, 
ceylonicus, T, fumifer, T, griseipalpis, T. hilaris, T. immanis, 
T. latifacies, T. malayensis, T. minimus, T. nemocallosuSy T. 
nemoralis, T. partitus, T. rubidujs, T. rufiventris, T, striatuSy T, 
vagus, T. vanecki, and T. tropicus. 


Observations on the monthly prevalence of tabanid flies were 
made. During a period of a little over 3 years, 854 flies were 
caught, of which 480 were Tabanus striatum and 374 Tabanus 
redueens. The male — ^female ratio of the former is 1 :7.42, the 
latter, 1 :25.71. A large number of males were collected during 
February and March. The peak of fly-incidence occurred in 

108S2 2 

384 The Philippine Journal of Science ^^^^ 

October or two months after the heaviest rainfall in August. 
A subpeak was noted in February following an increase in the 
rainfall during December. 

Positive results were obtained in experiments on surra trans- 
mission using Tahanus reducens and Tabantis striuttis. Tabanus 
reducens appeared to be a more efficient vector than T, striatus, 
due probably to its bigger size. 


Anonymous. Transmission of surra (see T. annamnese and T, evansi, 
126, 158) Rev. App. Ent., Ser. B: Med. and Vet. 28 (1941) 179. 

Cross, H. E. and P. G. Patel. Camel surra. Punjab Dept. Agri. Vet. 
BuU. No. S (1922) 1-19. 

Frasee, H. and S. S. Symonds. Surra in Federated Malay States. Studies 
Inst. Med. Res., F. M. S., No. 9 (1908). Cited by C. M. Wenyon, 
Protozoology II (1926) ix-1563 pp. 

Kblser, R. a. Transmission of surra among animals of the equine species. 
Philip. Jour. Sd. 34 (1927) 115-141. 

Kroeber, O. Die Tabaniden der Philippinen. Archiv. fur Naturgeschiechte. 
Abt. A, Heft 1 (1924) 1-27. 

Leesb, a. S. Final report for 12 months ending 31st March 1914, British 
East Africa. Summarized: Rev. of Applied Entomology 2 (1914) 

Manreza, M. and 0. Mondonedo. Studies on Surra, III. A survey of the 
incidence of surra in the vicinity of the College of Agriculture 
with observations on numerical fluctuation of tabanid flies. Philip. 
Agric. 24 (1935) 111-125. 

Mitzmain, M. B. The mechanical transmission of surra by Tabanus 
striatus. U. S. Bureau of Agriculture Bull. No. 28 (1913) 3-11. 

Mitzmain, M. B. Collected studies on the insect transmission of T. evansi, 
Treas. Dept. U. S. P. H. S., Hyg. Lab. Bull. No. 94 (1914) 7-39, 
Summarized: Trop. Vet. Bull. 5 (1917) 10-14. 

NiESCHULZ, O. Zoologische Bijdragen tot het Surraprobleem. III. Over- 
brengingsproeven met Tabanus rubidus Wied., T, striatus Fabr., en 
Stomaxys calcitrans L. Nederl. — Ind. Blad. v. Diergeneesk. 38 (1926) 
255-299. Summarized: Trop. Vet. Bull. 14 (1926) 129. 

NiESCHULZ, 0. Zoologische Bijdragen tot het Surraprobleem. XI. Enkele 
proeven met Haematopota trucata Schuurm. Stekh., H. Irrorata Macq. 
en Tabanus brunnipes Sch. Stekh. Veeartsenijkundige Mededeeling. 
Dept. Van Landbouw Nijverheid en Handel. Ned. Indie. No. 61 (1927) 
13 pp. Summarized: Trop. Vet. Bull. 15 (1927) 129. 

NiESCHULZ, O. Verdere surra — overbrengingsproeven met enkele tabani- 
den — ^soorten op Sumatra. Veeartsenijkundige Med. Dept. Van Land- 
bouw Nijverheid en Handel Ned. Indie. No. 66, 59 pp. Summarized: 
Trop. Vet. Bull. 17 (1929) 41-42. 

Patton, W. S. and F. W. Gragg. A Textbook of Medical Entomology. 
London, Madras and Calcutta. Christian Literature Society for 
India. 1913. viii— 765 pp. 

'^7''* Yutuc: The Prevalence of Tabanid Flies 385 

ScHUURMAN Stekhoven, Jr., J. R. The tabanids of the Dutch East Indian 
Archipelago. Treubia 6 (1926) 1-551. 

Sergent, Ed. and Et. El debab, trypanosomiase des dromadaires de 
rAfrique du Nord. Ann. Inst. Pasteur, 19 (1905 b) 17. Cited by 
C. M. Wenyon 1926. 

Sergent, Ed. and Et. Etudes sur les trypanosomiase de Berberie en 1905. 
Ann. Inst. Pasteur 20 (1906 a) 665. Cited by C. M. Wenyon 1926. 


Plate 1 

Fig. 1. Female and male Tabarvus reducens, 
2. Female and male Tabanus striatus. 


XVTVVI Pm-V.\U-XVK OF 1\.\B.\\H» Fl }F.S. | 

fPHtfji', ioi iL Sri.. Vol.. 7s. Hi%, I. 



By Edwin Bingham Copeland 
Of the University of Calif omidy Berkeley 


This paper includes material based on work mostly performed 
from 1939 to 1941, and included in several papers originally 
submitted for publication in The Philippine Journal of Science. 
It is recast here, in conformity with classification and nomen- 
clature of my Genera Filicum. When war made the publica- 
tion of these papers impossible, even after proof had been 
corrected, and the fate of my manuscripts unknown (all are 
known now to have been destroyed), I guarded the names of 
the many novelties by publication of diagnoses in Univ. Calif. 
Publ. Bot. 18 (1942) 217-226. The descriptions and illustra- 
tions, necessary to make these novelties certainly and easily 
recognizable, appear here for the first time. 

Key to Genera of New Guinea Aspidiaceae 

Fronds dimorphic. 
Veins free. 

Not widely scandent, fronds simple 14. Elaj}kofflos8um 

Rhizome scandent, fronds compound. 

Sori distinct 10. Thysanosoria 

Sporangia covering fertile surface. 

Terminal leaflet not articulate 9. Lomariopsis 

Distal leaflet jointed or suppressed. 

Leaves in two rows on rhizome 11. TeratophyUum 

Stipes polyseriate 12. Arthrobotrya 

Veins anastomosing. 
Without included veinlets. 

Widely scandent 13. Lomagramma 

Not scandent. 

Bhizome short, fronds clustered 20. Stenosemia 

Ehizome creeping, fronds remote 8. Bolbitis 

Included veinlets present. 

Bhizome short, fronds clustered 22. Hemigramma 

Bhizome creeping 8. Bolbitis 

* This paper is in part a report on the ferns collected by the third 
Archbold Expedition to New Guinea, in collaboration with the Government 
of the Netherlands Indies. Previous reports were published in the Philip. 
Jour. Sci. 73 (1940) 345 and 457; 75 (1941) 347; and 76 (1941) 23; 
and Univ. Calif. Publ. Bot. IS (1942) 217. 


390 The Philippine Journal of Science ^^^^ 

Fronds not dimorphic or slightly so. 
Veins free. 

Indusium wanting. 

Sori round 23. Lastrea 

Sori elongate 24. Currania 

Indusium elongate along vein. 

Pinnules articulate, subdimidiate 6. Didymochlaena 

Indusium ovate, free at apex 2. Stenolepia 

Indusium round, bursting on top 1. Diaealpe 

Indusium peltate. 

Pinnae articulate to rachis 5. Cyclopeltis 

Pinnae not articulate. 
Articulate hairs wanting. 

Frond dilated at base 7. Rumohra 

Frond not widened at base 4. Polystichum 

Articulate hairs on rachis 17. Dryopolystichum 

Indusium renif orm. 

Nodes of rachis enlarged 8. Acrophorus 

Nodes not enlarged. 

Axes bearing pluricellular hairs 16. Ctenitis 

Glabrous, or hairs unicellular. 

Each sinus occupied by a tooth 18. Pteridrys 

Sinuses not tooth-bearing. 

Minor axes decurrent on major 15. Dryopteris 

Minor axes not decurrent. 

Frond anadromic, base broad 7. Rumohra 

Frond catadromic 23. Lastrea 

Fronds anastomosing in regular pairs. 
Frond not proliferous. 
Bachis not spiny. 

Indusium roundish, fixed by sinus 25. Cyclosorus 

Indusium oblong, fixed along middle 27. Sphaerostephanos 

Bachis spiny, sori linear 30. Callipteris 

Frond proliferous 26. Ampelopteris 

Veins anastomosing irregularly. 

Areolae costular only, costal sori elongate 19. Heterogonium 

Areolae not all costular, or sori round. 

Free included veinlets present 21. Tectaria 

Free included veinlets wanting. 

Indusium opening along margin 28. Athyrium 

Indusium bursting on top 29. Diplaziopsis 

1. DIACALPE Blume 


D, aspidioides Blume, Enum. (1828) 241. 

Carr. U567, Boridi, alt. 1,800 m; Brass Ui5, Mt. Tafa, alt. 
2,300 m. 
To India and China. 

'''** Copeland: Aspidiacme of New Guinea 391 

2. STENOLAPIA Van Alderwerelt Van Rosenburgh 

S. TBISTIS (Bliiine) vJi.v.R. 

S. tristis (Blume) v.A.v.R., Bull. Dept. Agric. Ind. N^rl. No. 27 (1909) 
46, PI. 7. 

var. paptmna C. Chr., Brittonia 2 (1937) 294. 
var. reducta C. Chr., ibid. 

Brass U2SU, iS29, US2, Mt. Albert Edward, alt. 3,680 m; 
Clemens, Mt. Sarawaket. 

The species is known elsewhere to be very plastic under the 
influence of exposure, and var. reducta is a stunted form. 

The species in Java and Borneo. 


A. STIPELLATUS (WaUich) Moore 

A. stipellatus (Wallich) Moore, Gard. Chron. (1854) 135. 

A. stipellatus var. montana Ros., Pedde's Repert. 12 (1913) 166. 

A. Blumei Ching, Gardens' Bull. 7 (1934) 226. 

Brass 10716, Lake Habbema, alt. 2,800 m; 12219, alt. 1,750 
m; Keysser 11 7S, II 76 (isotype of var. montana), Bolan, alt. 
2,400-3,000 m. 

Paying some regard to geographic distribution, and judging 
by specimens I have seen, alive and in herbaria, I feel constrained 
to construe Acrophorus as a genus of one known variable species. 
It varies in size, and therewith it may be three times or four 
times pinnate. It varies in size of pinnules, in prickliness, 
chaffiness and hairiness of the axes, and in size of indusia. But 
there is not an Indian species characterized by one set of char- 
acters, nor a Javan species with another set, which might be 
the A. stipellatus and A. Blumei of Ching and Christensen. 
Wallich 359, from Nepal, is fully quadripinnate, with very 
small ultimate pinnules; its axes are v^ry nearly smooth, and 
the indusia small; but specimens like it in the four characters 
mentioned are in hand from Sumatra and Fiji. Borneo and 
Luzon specimens are most scaly, and this strain extends to 
Formosa. Judging by material in hand, the species runs small 
in China, and very large in Fiji, 


Acrophorus loxoseaphoides (Baker) Alston, Jour, of Bot. (1939) 288. 
Polypodium loxoseaphoides Baker, Jour, of Bot. 2S (1890) 107. 
Dryopteris loxoseaphoides C. Chr., Index (1905) 276. 
Leucostegia loxoseaphoides C. Chr., Suppl. Ill (1934) 120. 

392 rfee Philippine Journal of Science 1^49 

"The type is an exceedingly poor specimem, but this appears 
to be the correct disposition of it," — ^Alston, 1. c. I have no 
other opinion, but would emphasize the word appears. The 
type was collected by MacGregor, "Mount Musgrave and 9,200 
ft. on Owen Stanley range." 

Dryopteris subdigitata Brause, Engler's Jahrb. 56 (1920) 94, 
was also treated as Leucostegia by Christensen, SuppL III 98 
and 121, with the suggestion that it, and Davallia Forbesii 
Gepp, Journal of Bot. (1923) Suppl. 59, might both be Leiuws- 
tegia hxoscaphoides. 


Key to the species 

Frond over half as broad as long 1. P, keysserianum 

Frond over twice as long as broad. 
Pinnules or segments not strongly bullate. 
Lamina at least 30 cm long. 

Freely tripinnate 2. P. bamlerianum 

Only lowest pinnules or none pinnate. 
Pinnnles acute or aculeate. 

Segments of pinnae mostly connected 3. P. aculeatum 

Pinnules mostly free. 

Free pinnules less than 15 pairs. 
Sori costal to medial. 

Some pinnae over 15 cm long 3. P. aculeatum 

Pinnae smaller. 

Pinnules weakly mucronate 3. P. aculeatum 

Pinnules very mucronate 4. P. Archboldii 

Sori submarginal 5. P. Brassii 

Free pinnules about 20 pairs 6. P. bolanicum 

Pinnules rounded, hardly mucronate. 

Pinnules crenate to entire 7. jp. muticum 

Pinnules incised to subpinnate 8. P. Myer-Dreesii 

Lamina usually under 25 cm long. 

Stipes conspicuously gtout 9. P. papuanum 

Stipes slender. 

Pinnules acute 10. P. kinabaluense 

Pinnules rounded 11. P. alpinum 

Pinnules and segments strongly bullate. 

Frond lanceolate 12. P. cheilanthaides 

Frond linear 13. P. limare 


P. keysserianum Ros., Fade's Repert. la (1912) 331. 

Keysser 60, Sattelberg, alt. 1,600 m, isotype in Herb. Univ. 
Calif.; also, Keysser 19, much less dissected than the tjrpe. 

'^'^'^ Copeland: Aspidiaeeae of New Guinea 393 


P. bamlerianum Ros., Fedde's Repert. 10 (1912) 330. 

Bamler K 10, Cromwell Mountains, alt. 1,600 m, isotype in 
Herb. Univ. Calif. Our specimen of Bamler K 10 a, var. 
ferrugineo-paleacea, a segregate from the type collection, is also 
tripinnate, but is more ample and more scaly ; it is approximately 
P. moluccense (Blume) Moore; but as it does not fit Rosenstock's 
varietal description, some error is suspected. Endemic. 

3. P. ACUL£ATUIII (L.) Schott 

P. aculeoitum (L.) Schott, Genera (1834) PI. 9, note. 

Construed in its usual broad sense, this can include Brass 
12666, alt. 2,150 m; 13665, alt. 700 m; and 1S38A, alt 850 m; 
Bamler 188 (1914) ; and Keysser 208; these represent a single 
species. Related to it is Schlechter 16921, received as P. 
aeuleatum var. suhamoenum Christ, a doubtful identification. 
Keysser 201 is distinct, but is more like the P. aeuleatum of 
Europe than is any other New Guinea collection. 

Cosmopolitan, as construed. 

4. P. ARCHBOIiDII Copel. Plate 1. 

P. Archboldii Copel., Univ. Calif. Publ. Bot 18 (1942) 221. 

P. gregis p. aculeati lamina coriacea rhachi inter alias paleis 
lanceolatis patentibus onusta; rhizomate adscendente 5-8 mm 
crasso, paleis atrocastaneis 1 cm longis basibusque stipitum 
profunde immerso; stipite 30 cm alto, erecto, sordide fusco, 
paleis variis maximis 13 mm longis 2.5 mm latis apice acicula- 
ribus dentibus minutis remotis recte distantibus ciliolatis omato 
et furfuraceo; lamina 40 cm longa, 12 cm lata, acuminata, 
subtripinnata, rhachi paleis 5 mm longis lanceolatis aciculari- 
acuminatis rectis vel tortis integris vel ciliatis ornata aliisque 
variis et acicularibus et amorphis dense vestita; pinnis inflmis 
8 cm longis vix deflexis, sequentibus 9 cm longis breviter (2 mm) 
pedicellatis, acutis, basi oblique truncatis, rhachibus paleis 
minoribus vestitis; pinnulis ca. 15-paribus, mucronatis, infimis 
acroscopicis maximis ca, 17 mm longis, basi obliquis, pinnula 
secundaria una acroscopica, pinnulis sequentibus auriculatis 
gradatim apicem brevem incisum versus pinnae decrescentibus, 
utraque facie pilis paucis debilibus vestitis ; soris inf ramedialibus, 
indusiis parvis peltatis f ugacibus. 

Dutch New Guinea: 7 km N-E. of Wilhelmina-top, alt. 3,560 
m. Brass & Myer-Drees No. 9850. "Subalpine forest ; occasional 
on mossy forest floor; clumps 60-80 cm high; fronds 3 or 4." 

394 The Philippine Journal of Science ^^^^ 

5. F. BEASSn Copel. sp. nov. Plate 2. 

P, gregis P. aculeati tenuiter coriaceum soris submarginalibus ; 
rhizomate ignoto; stipite 30 cm alto, gracile, fusco, deorsum 
paleis nigro-f uscis 5 mm longis lanceolatis acuminatis appressis 
vel inflexis sat dense minute denticulato-ciliatis, sursum paleis 
setiformibus haud dense vestito; lamina 30 cm longa, 12 cm lata, 
abrupte acuminata, apice 8 cm longo tantum pinnato, alibi 
bipinnata, rhachi atropurpureo-fusca paleis 1.5-3 mm longis 
linearibus contortis castaneis basibus dilatatis hinc inde rami- 
feris strigosa; pinnis infimis 7 cm longis subdeflexis, sequentibus 
8 cm longis falcato-adscendentibus, acuminatis, breviter (1.5 
mm) pedicellatis, rhachibus deorsum setoso-paleolatis sursum 
nudis; pinnulis infra apicem inciso-pinnatifidum pinnae ca. 12, 
mucronatis, infima acroscopica maxima 1 cm longa, 5-6 mm lata 
incisa, sequentibus gradatim decrescentibus contiguis interdum 
oblique rhomboideis argute serratis dentibus minute mucronatis, 
utraque facie mox glabris, superne f uscis inferne pallide oliva- 
ceis ; soris submarginalibus parvis, indusio minuto peltato caduco. 

Dutch New Guinea: Bele River, alt. 2,800 m; Brass No. 10247. 
"Occasional tufts in forest.'' 


P. bolamcum Kos., Fedde's Repert. 12 (1913) 170. 

Bolan, alt. 3,400-3,800 m, Keysser B 8 1912, isotype in Herb. 
Univ. Calif. Perhaps too near to P. microphyllum (Blume) 


Bolan, alt. 2,400-3,000 m, Keysser B 42, also in Herb. Univ. 
Calif., tearing the comment "potius spec, propria,'' with which 
I agree, although the species and variety are evidently nearly 
related. Endemic. 

7. p. MUnCUM Copel. Plate 3. 

P. muticum Copel, Univ. Calif. Publ. Bot. IS (1942) 221. 

Rhizomate erecto, paleis fuscis 2-2.5 cm longis linearibus 
aciculatis integris dense immerso; stipitibus fasciculatis, 15-18 
cm altis, 1.5 mm crassis, ochroleucis, paleis patentibus concolo- 
ribus medio brunnescentibus 5 mm longis ovatis oblique acutis 
nonnulis mucronulatis papyraceis primo integris denique plus 
minus laceris postquam dejectis cicatrices fuscas relinquentibus 
omatis; lamina 35 cm longa, 13 cm lata, acuminata, bipinnata, 
rhachi quam stipite densius paleis minoribus vestita; pinnis 

^"^'^ Copeland: Aspidiaceae of New Guinea 395 

infimis 5 cm longis 2 cm latis defl'exis, medialibus 7 cm longis 
basi 2.5 cm alibi 2 cm latis, subfalcatis, obtusis vel breviter 
mucronulatis, breviter pedicellatis, rhachibus paleolatis sursum 
nudis; pinnulis ca. 9-paribus, contiguis, oblique oblongis, rotun- 
datis, undulato-crenatis vel fere integris, subcoriaceis, glabris; 
soris medialibus, e facie superiore laminae expressis, indusio 
parvo orbiculare sat persistente. 

Dutch New Guinea: Bele River, alt. 2,200 m; Brass lUS^, 
type, "Numerous clumps on dry face of limestone cliff in forest 
shade;" No. 11^38, from the same place, a small juvenile plant, 
with particularly small paleae, the pinnae with 0-2 basal 

8. P. MYER-DREESU Copel. Plate 4. 

P. Myer-Dreesii Copel., Univ. Calif. Publ. Bot. 18 (1942) 221. 

P. mutico affine, paleis rhizomatis atrocastaneis, stipitis 
rhacheosque castaneis aciculari-lanceolatis, pinnulis pauUo 
majoribus, infimis basi pinnula secundaria una prseditis, 
sequentibus pinnatisectis, indusiis magnis interdum excentrice 
adnatis etenim rarissime orbiculari-reniformibus diversum. 

Dutch New Guinea: 7 km N-E. of Wilhelmina-top, alt. 3,560 
m. Brass & Myer-Drees No, 10033; "Clumped under rocks in 
subalpine forest.'' 

9. P. PAPUABOnU C. Chr. 

P. papuanum C. Chr., Brittonia 2 (1937) 299. 

Brass 9136, Lake Habbema, alt. 3,225 m, "Forming dense 
clumps 20-30 cm high and up to 1 m in diameter in forest 
glades;" Brass & Myer-Drees 10055 and 10112, Mt. Wilhelmina, 
alt. 3,950 m. No. 10055 is "one of the few plants on limestone 
screes; clumps 20-40 cm high, scattered amongst the rocks." 
It and No. 9136 are more or less dwarfed. No. 10112, "Under 
a rock on Alpine grassland," has responded to its more sheltered 
position by producing somewhat more ample fronds, almost flat 
pinnules, and longer, paler, less congested paleae on stipe and 
rachis. The previous collection are Brass 4350 (type) and 
4^690, dwarfed. 

10. p. KINABALUENSE C. Chr. 

P. Jdnabalmme C. Chr., Gardens* BuU. 7 (1934) 255. 

Mt. Albert Edward, alt. 3,680 m, Brass 5678, det. Christensen. 

396 The Philippine Journal of Science 1^49 

11. p. ALPINUM Bf>s. 

P. alpinum Ros., Pedde's Repert. 12 (1913) 171, 

Brass H83, Lake Habbema, alt. 3,225 m, "under rocks in 
edge of forest; fronds spreading"; Brass & Myer-Drees 9766, 
9967, 10838, Mt. Wilhelmina, alt. 3,500-3,800 m. Variable, and 
none exactly like the type, Keysser B 5 (1912), isotype in 
Herb. Univ. Calif. Judging by its long and comparatively 
naked stipe, the type grew in a sheltered place, perhaps under 
a rock. Endemic. 

12. p. CHEILANTHOmiS Copel. Pl«te 5. 

P. cheilanthoides Copel., Univ. Calif. Publ. Bot. IS (1942) 221, 

Papuapteris ampla subtripinnata, rhizomate erecto, 2 cm (cum 
tesibus stipitum, 4 cm) crasso, basibusque stipitum paleis 
primo rufis, deinde atro-f uscis tum demum nigris nitidis usque 
ad 3 cm longis 3 mm latis aculeato-attenuatis primo integris 
demum fisso-ciliatis vestitis; stipitibus fasciculatis, 25-30 cm 
altis 3-5 mm crassis, paleis sursum decrescentibus et variis 
densis vestitis etiam squamulosis; lamina ca. 35 cm longa, 5-7 
cm lata, rhachibus paleis polymorphis et lanceolatis majoribus et 
ovatis minutis et setiformibus plerisque strigoso-ciliatis sursum 
pall^centibus ubique dense onustis; pinnis infimis ca. 4 cm 
remotis pauUo reductis hand deflexis, medialibus 5-6 cm longis 
1 cm latis, adscendentibus, sessilibus, obtusis; pinnulis infimis 
usque ad 9 mm longis, pinnatis cum pinnulis secundariis 1- 
3-paribus, sequentibus rarius pinnatis saepius pinnatifidis, ple- 
risque integris oblongis rotundatis buUatis, coriaceis, supeme 
plus minus decidue crinito-setosis, inferne dense paleatis et 
strigosis; soris infra paleas et margines revolutos plus minus 
occultis, indusiis ut videtur nullis, cellulis annulorum ca. 17, 
sporis oblongis densissime echinatis. 

Dutch New Guinea: 11 km N-E. of Wilhelmina-top, alt. 3,400 
m; Brass & Myer-Drees No. 9801, type, "Grassy bottom of a 
stream; fronds erect in large clumps;" also. No. 10102, alt. 
4,100 m. "Common on rocky grassy slopes ; clumps 60 cm high ; 
southern slopes of Mt. Wilhelmina." 

I believe that most pteridolc^ists will agree in placing this 
species in Polystichum. In the Orient, it is rather isolated by 
its strongly buUate pinnules, but this would not be so among 
Andean species of the genus. 

'■^'^ Copeland: Aspidiaceae of New Guinea 397 

13. p. LINEARE (C. Chr.) Copel., comb. hov. 

Papuapteris linearis C. Chr., Brittonia 2 (1937) 300, /. 2. 

Brass 4S0i, Mt. Albert Edward, alt. 3,690-3,980 m; cited 
also from Owen Stanley Range and Mt. Victoria. 

Endemic, and apparently of endemic parentage. Nearly 
related to the preceding species. 


Key to the species 

Pinnae not hastate 1. C. presliarui 

Pinnae hastate on upper side 2. C novoguineensis 

1. C. PRESLIANA (J. Smith) Berkeley 

C, presliana (J. Smith) Berkeley, Crypt. Bot. (1857) 517. 

Brass 8860, Hollandia, sea level; Schlechter 16284', alt 300 m; 
Bamler 64 (1909), Logaueng; King 469, Papua. 

Malaya; Philippines. Specimens from the Caroline and Gil- 
bert Islands and Tonkin have been referred to this species. 


C novoguineensis Ros., Fedde*s Repert. 10 (1912) 329. 

Known locally only by the type collection, Bamler, Rosen- 
stock FiL novog. exsicc. n. 86, Logaueng, alt. 300 m; isotype 
in Herb. Univ. Calif. 

Solomon Islands. 

D. TRUNCATULA (Swartz) J. Smith 

Brass 12155, alt. 1,750 m ; Schlechter 17694, alt. 300 m ; Bamler, 
Rosenstock Fil novog. exsicc. n. 225, Sattelberg, alt. 900 m; 
King 484, Papua, All New Guinea specimens represent var. 
ozeanica Ros., Fedde's Repert. 5 (1908) 374, characterized 
chiefly by small pinnules. 

7. RUMOHRA Raddi 

Key to the species 

Indusium large, peltate 1. R. adiantiformis 

Indusium small, round-reniform to peltate 2. R, anstata 

Indusium wanting 3. R, Hasseltii 

1. R. ADIANTIFORMIS (Forster) thing 

R. adiantiformis (Forster) Ching, Sinensia 5 (1934) 70. 

Dryopteris discophora Ros., Fedde's Repert. 12 (1913) 172. — 
"Eine durch ihre grossen schildformigen Indusien sehr auffal- 

398 The Philippine Journal of Science ^^^® 

lende Art." Isotype in Herb. Univ. Calif.; Clemens s. n., Mt. 
Sarawaket, alt. 3,000-10,000 feet. 
All Southern lands. 

2. R. ARISTATA (Forster) Ching 

R. wriatata (Forster) Ching, Sinensia 5 (1934) 50. 

Brass 5177 and 5202, Bamler 49p and Bamler, Rosenstock Fil. 
novog. exsicc. n, 166 are this species, broadly construed. Carr 
14388 is similar but distinct, but does not well fit Blume's 
description of Aspidium appendiculatum. 

India across Polynesia. 

3. R. HASSELTn (Blnme) Ching 

R. Hasseltii (Blume) Ching, Sinensia 5 (1934) 61. 

Bamler 110 (1914), Sattelberg, alt. 800-900 m. 
Java to Leyte, Formosa and Assam. 

S. BOLBinS Schott 

Genera (1834) Plate U; Ching, in C. Chr., Suppl. Ill (1934) 47. 
Campium PresL; Copel., Philip. Jour. Sci. 37 (1928) 341 partim. 

Because the formation of fertile fronds is seasonal or occa- 
sional, this genus is likely to be passed over by collectors. It 
seems to be poorly developed in New Guinea, where only 5 
species have been reported, in contrast with 15 in the Philip- 

Key to the species 

Apex of frond a leaflet, like lateral pinnae 1. B, heteroclita 

Apex formed by fusion of decrescent pinnae. 
Pinnae shallowly cut. 

Lamina over 30 cm long 2. B. quoyana 

Lamina smaller. 

Main veins evident 3. B. Taylori 

Main veins undeveloped 4. B. parva 

Pinnae deeply pinnatifid 5. B, a/rguta 

1. B. HETEROCLITA (Presl) Ching 

B. heteroclita (Presl) Ching, in C. Chr., Suppl. Ill (1934) 48. 
Campium heteroclitum Copel., Philip. Jour. Sci. 37 (1928) 359, /. IS, 

Idenburg River, alt. 250 m. Brass 18242; Papua, King 265, 
Caroline Islands to China and India. 

2. B. QUOTANA (Gaud.) Ching 

B. quoyana (Gaud.) Ching, ibid. 49. 
Campium quovanum Copel., ibid, 366, /. 20, 

'^'^*^ CopeUmd: Aspidiaceae of New Guinea 399 

Schlechter 16236, Kaiser-Wilhelmsland. From Papua, I have 
four collections by King, all showing a tendency to confine the 
sporangia to a marginal band, and leave the costal area of 
fertile pinnae sterile. One of these was named Leptochilus 
cttspidattis (Presl) var., marginalis Ros., Fedde's Repert. 9 
(1911) 426. They also represent Heteroneuron Naumanni 
Kuhn, described from New Hanover, which I do not regard 
as distinct. 

New Hebrides to Java and Samar. 

3. B. TATLORI (Bailey) Chlng 

B. Taylori (Bailey) Ching:, ibid. 50. 
Campium Taylori Copel., ibid. 374. 

Brass 128SS, "abundant on rocks in a rain-forest stream at 
1,150 m alt," 6 km SW. of Bernhard Camp, Idenburg River, 
is approximately this species. Queensland. 

4. B. PARVA (Copel.) Ching 

B, pa/rva (Copel.) Ching, ibid. 49. 

Campium parvum (Copel., ibid. 375, PI. 21 and /. 28. 

Schlechter 16168, Kaiser-Wilhelmsland, alt. 300 m. King 269 
from Papua, is similar in appearance. A single specimen from 
Mrs. Clemens, from Morobe, alt. 4,200 feet, looks very distinct, 
being larger, with few broad pinnae, but may possibly be the 
same species in ample development. 


5. B. ARGUTA (F^e) Ching 

B. arguta (Fee) Ching, ibid. 47. 

Heteroneuron argutum F4e, Acrost. (1845) 96, PI. 25, f. 2. 

Campium argutum Copel., ibid. 376, PI. 22. 

Brass lSi7S, "Several tufts on the bank of a rain-forest 
stream at 800 m alt," 2 km SW. of Bernhard Camp, Idenburg 


In the Philippines, where they are better known, some Bolbitis 
species are notoriously variable. With the three preceding 
species represented by a total of 5 collections, of which no two 
are fully identical, their distinctness and their identity are for 
the present uncertain. 

10882 S 

400 The Philippine Journal of Science ^^^^ 


Key to the species 

Fertile pinnae up to 3 mm wide. 

Sterile pinnae lanceolate 1. L. coehinchinensis 

Sterile pinnae oblanceolate 2. L. Kingii 

Fertile pinnae 5-10 mm wide 3. L. subtrifoliata 

Fertile pinnae up to 20 mm wide 4, L. intermedia 


L. coehinchinensis Fee, Acrost. (1845) 68, PL 27; Holttum, Gardens' 
Bull. 5 (1932) 266. 

Bamler 50, Bamler, Rosenstock, Fil. novog. exsicc. n, 251 y 
Sattelberg, alt. 300-700 m. 
To Indo-China. 

2. L. KINGn (Copel.) Holttum 

L. Kingii (Copel.) Holttum, Gardens' Bull. 5 (1932) 273. 
Stenochlaena Kingii Copel., Philip. Jour. Sci. 6C (1911) 80. 

Known only by the type collection, King 285, from Papua; 
perhaps not sufficiently distinct from L. coehinchinensis. 

3. L. SUBTRIFOLIATA (Copel.) Holttum 

L. subtrifoliata (Copel.) Holttum I.e. 274. 

Brass 13882, alt. 120 m, climbing 4-5 m on small trees. 
Typical except for fewer pinnae, larger sterile pinnae, and less 
scaly stipe; pinnae sessile. 

Southern Philippines. 

4. L. INTERMEDIA (Copel.) Holttum 

L. intermedia (Copel.) Holttum, 1. c. 
Stenochlaena intermedia Copel., Philip. Jour. Sci. 7C (1912) 67. 

Known only by the type collection, King 870, from Ambasi, 
Papua. Very distinct. 


T. PTERIDIFORMIS (Cesati) C. Chr. 

T. pteridiformis (Cesati) C. Chr., Suppl. Ill (1934) 187; Dansk 

Bot. Arkiv 9 No. 3 (1937) 51. 
T. dimorphophylla Gepp, Dutch N. W. New Guinea (1917) 193, Plate 4. 

Arfak Peninsula, Cesati, Gihhs. Unknown to me. 

11. TEBATOPHYLLUM Mettenius 
T. ACULEATUM (Blume) Mett. 

r. aculeatum (Blume) Mett., in Kuhn, Ann. Mus. Lugd Bat. 4 (1869) 
296; Holttum, Gardens' Bull. 5 (1932) 284, Pis. 2, S, and figs. SS-AO. 

'^'^»* Copeland: Aspidiaceae of New Guinea 401 

Werner 6i, Damun; also cited by Holttum, 1. c. 289, are Lam 
h2U and 1105. 
Malaya; Philippines. 


A. articulata J. Smith, Hist. Fil. (1875) 142. 

Brass 12202, Idenburg River, alt. 1,700 m; Clemens 11011, 
Morobe; King i90, Brass 1468, Papua. 
Philippines; Celebes; Solomon Islands. 

13. LOMAGRAMMA J. Smith 

Key to the species 

Sterile pinnae at least 15 mm wide. 

Pinnae with winged stalks 1. L, sinuata 

Pinnae hardly stalked 2. L. novoguineensis 

Sterile pinnae 8 mm wide 8, L. angustipinna 

1. L. SINUATA C. Chr. 

L. sinuata C. Chr., f. papttana C. Chr., Brittonia 2 (1937) 302. 

Brass 187 52, Idenburg River, alt. 75 m; scandent 7-8 m. 
This is a perfect match for Brass 5195, the basis of the forma. 
It is to be observed that the very large pinnae are short-stalked. 
Holttum, Gardens' Bull. 9 (1937) 216 has noted that the long 
pedicels of the pinnae, originally a characteristic of the species, 
are inconstant. The fertile pinnae are 3-4 mm wide. 

The species in New Guinea, Celebes, Borneo and Java. 

2a. L. LOMARIOIDES (Blume) J. Smith? 

L. novoguineensis (Brause) C. Chr., Suppl. Ill (1934) 124. 
Leptochilus novoguineensis Brause, Engler's Jahrb. 56 (1920) 117. 

Ledermann 9524, alt. 850 m; Docters van Leeuwen 9616, 
Mamberamo River, alt. 50 m. 
Endemic. See Holttum, Gardens' Bull. 9 (1937) 209, PMe 15. 

2a. L. LOMARIOIDES (Blume) J. Smith? 

Brass 13751, with L. sinuata, scandent to 8 m. Pinnae sessile, 
broadly obliquely cuneate, rather thin, but not satisfactorily 
distinguishable from the Java plant; fertile pinnae about 2 mm 

Brass 12950, alt. 1,200 m, scandent 3-4 m. Pinnae sessile 
with broad, approximately truncate bases ; fertile pinnae 4 mm 
wide; stipes 10 cm long. Except for the long stipe, this would 
be more like L. BrooksiL 

402 The Philippine Journal of Science ^^^^ 

These two specimens are conspecific, whatever the species ; and 
I suspect that they are variants of a species including No. 13752. 
Altogether, there are too few and too imperfect collections in 
this genus to afford dependable conclusions as to variability. 
L. lomarioides is regarded by Holttum as strictly Javanese. 

3. L. ANGUSTIPINNA Copel. Plate 6a. 

L, angmtipinna Copel., Univ. Calif. Publ. Bot. 18 (1942) 222. 
Rhizomate scandente, sicco 5 mm crasso, basique stipitis paleis 
atrocastaneis 1-2 mm longis late lanceolatis squamulisque 
amorphis brunneis sparsis; bathyphyllis ignotis; frondis sterilis 
stipite 14 cm longo, gracile, stramineo; lamina 50 cm longa, 
20 cm lata, pinna terminate 7 cm longa non articulata, pinnis 
lateralibus ca. 30-paribus articulatis, inf erioribus decrescentibus, 
medialibus ca. 10.5 cm longis 8 mm latis, falcato-acuminatis, 
subsessilibus, basi basiscopice angustissime acroscopice latius 
cuneatis, integris, herbaceis, costa inf erne decidue paleolatis; 
venis tenuibus inconspicuis, areolarum seriebus 2 vel 3 irregula- 
ribus; rhachi paleis fuscis linearibus ca. 1 mm longis basi 
interdum dilatatis sparsa ; frondis fertilis pinnis breviter (2 mm) 
stipitulatis, usque ad 9 cm longis et 3 mm latis, sporangiis 

Dutch New Guinea: 2 km SW. of Bernhard Camp, Idenburg 
River, alt. 750 m, Brass 1SM6; "Climbing epiphyte abundant 
on substage trees." Type in Gray Herbarium. 

In spite of doubt as to the stability of various characters, 
and as to the distinctness of species based on them in this 
genus, I find it impossible to refer this specimen even appro- 
ximately to any known species. The narrow sterile pinnae 
are unique in the genus, as is also the combination of thin 
lamina and inconspicuous venation. The dark-reddish sporan- 
gia suggest those occasionally found on more than one species 
of Bolbitis, and on Teratophyllum WilliamsiL 


Key to the species 

Small plants, lamina under 5 cm long. 
Paleae light-brown, subciliate. 

Stipe of sterile frond under 1 cm long 1. E, bolanicum 

Stipe of sterile frond over 2 cm long 2. E, habbemense 

Paleae dark, ciliate 8. E, helhmgianum 

'''^'^ Copeland: Aspidiaceae of New Guinea 403 

Larger plants. 
Fronds clustered or approximate. 

Fronds narrowly lanceolate, scaly 4. E, petiolatum 

Fronds broader or glabrescent. 

Margin conspicuously fringed 5. E, Cumingii 

Margin not fringed. 

Stipe conspicuously scaly 6. E. sordidum 

Stipe glabrescent. 
Lamina conspicuously decurrent. 

Lamina under 30 cm long 7. E.novoguineense 

Lamina over 40 cm long 8. E. Archboldii 

Lamina acute at base 9. E, sclerophyllum 

Bhizome elongate, fronds seriate. 

Lamina half as wide as long 10. E, laticuneatum 

Lamina relatively narrow. 

Paleae broad, light-brown 11. E, angulatum 

Paleae narrow and dark. 

Base of lamina abruptly narrowed 12. E, brunneum 

Base cuneate or decurrent. 

Rhizome 1.5-2 mm thick 13. E, repens 

Rhizome much stouter 14. E, fuscum 


E. bolanicum Ros., Fedde's Repert. 12 (1913) 180. 

Bolan Mte. alt. 2,400-3,00(3 m, Keysser (1912) B 62; iso- 
type in Herb. Univ. Calif., including sterile frond. 

2. £. HABBEMENSE Copel. Plate 7. 

E. habbemense Copel., Univ. Calif. Publ. Bot. 18 (1942) 226. 

Rhizomate late repente, 1-2 mm crasso, paleis ferrugineis 
1-2 mm longis plerisque ovatis subintegris haud dense vestito; 
stipitibus ca. 1 cm remotis, frondis sterilis ca. 2.5 cm longis, 
1 mm crassis, exalatis, paleis iis rhizomatis conformibus sursum 
decrescentibus vestitis, frondis fertilis breviore; lamina sterile 
ca, 2.5 cm longa, 8 mm lata, utrinque acuta, mox glabrescente, 
rigide coriacea, fusca, margine cartilagineo subdeflexo; lamina 
fertile conf orme sed basi rotundata. 

Dutch New Guinea: Lake Habbema, alt. 3,225 m, Brass No. 
9083, type in Gray Herbarium. "In a cushion of hepatics on 
exposed branch of tree.*' Only one plant, with one fertile and 
several sterile fronds. 

A dwarf, perhaps related to E, laurifoUum, but not to the 
other dwarfs known in New Guinea. 


JS'. hellwigianum Bos., Nova Guinea 8 (1912) 731. 

404 The Philippine Journal of Science ^^^^ 

Dutch New Guinea: summit of Hellwig Mts., von Roemer 
127S. Not seen. 

4. E. PEnOLATUM (Swartz) Urban 

E. petiolatum (Swartz) Urban, Symb. Ant. IV (1908) 61. 

Keysser (1910) K 11; Brass 5^8^, Mafulu, alt. 1,250 m. I 
have in hand both of these collections, both determined as 
E, Copelandii Christ. Neither exactly matches that Philippine 
species, which is a member of the world-wide group of E, 
petiolatum. As an alternative name for the New Guinea plant, 
I would prefer E. Blumei J. Smith. 

5. E. CUMINGn (F6e) Moore var. PAPUANUM C. Chr. 

E, Cumingii (Fee) Moore, var. papuanum C. Chr., Brittonia 2 (1937) 

Mt. Tafa, alt. 2,300 m. Brass U12. 

This is a satisfactory match for some Philipppine specimens. 

6. E. SORDIDUM Christ 

E. sordidum Christ, Nova Guinea S (1909) 156. 

Dutch New Guinea: North River, near Geitenkamp, Versteeg 
HS2. Not seen. Brause lists 5 subsequent collections by 
Ledermann in Kaiser-Wilhelmsland. Christ remarked that this 
was the first Elaphoglossum known in New Guinea. 


E, novoguineense Eos., Fedde's Eepert. 10 (1912) 341, 
E. Brassii C. Chr., Brittonia 2 (1937) 316. 

Sattelberg, Bander 67 (1909), isotype in Herb. Univ. Calif.; 
ihid., Keysser 226 (1913) ; Mafulu, alt. 1,950 m. Brass 5558, 
E. Brassii; Bele River, Brass 1H98, 


8. £. ARCHBOLDU Copel. Plate 8. 

E. Archboldii Copel., Univ. Calif. Publ. Bot. 18 (1942) 226. 

Rhizomate breve, crasso, stipitibus radicibusque ubique obsito, 
etiam paleis sordide fuscis ca. 2 mm longis ovatis integris 
vestito; stipitibus fasciculatis, basibus (phyllopodiis) castaneis 
3-4 cm altis, 3-4 crassis, paleis quam iis rhizomatis longioribus 
sparsis; frondis sterilis stipite 10-30 cm alto, decurrenti-alato, 
fusco; lamina 40-50 cm longa, 5.5 cm lata, acuta, coriacea, 
margine subdeflexo, utraque facie glabrescente, venis manifestis, 
furcatis; frondis fertilis stipite 30-45 cm alto, sursum brevi- 

'^'^'^ Copeland: Aspidiaceae of New Guinea 405 

alato, glabrescente, lamina ca. 30 cm longa, 2-3 cm lata, costa 
deorsum et margine angusto exceptis sporangiis densissime 

Dutch New Guinea: 4 km SW. of Bernhard Camp, Idenburg 
River, alt. 850-900 m, Brass 13220, type in Gray Herbarium; 
ibidem, No. lSi8^. 

A large relative of E. callifolium, with smaller and darker 
paleae, the stipe of the sterile frond winged ahnost to the base. 


E. sclerophyllum v.A.v.R., Nova Guinea 14 (1924) 22. 
Doorman-top, alt. 3,200 m, Lam 1798. Not seen. 

10. E. LATICUNEATUM Copcl. Plate 9. 

E. laticuneatum Copel., Univ. Calif. Publ. Bot. IS (1942) 226. 

Rhizoma te late repente, 2-3 mm crasso, paleis fuscis nigres- 
centibus ovato-lanceolatis 2-3 mm longis acuminatis irregulariter 
ciliatis vestito; stipitibus valde remotis, phyllopodiis obscuris 
paleaceis 1-1.5 cm altis, frondium sterilium ca. 8 cm altis fusco- 
stramineis squamulis paucis onustis; lamina sterile 6-8 cm 
longa, 2.5-4 cm lata, apice rotundata, basi cuneata, coriacea, 
margine angustissime cartilaginea, utraque facie squamis 
elongato-triangularibus acuminatis 1 mm longis pauci-ciliatis 
atris sparsa; frondis fertilis stipite 15 cm alto, lamina 8 cm 
longa, vix 4 cm lata, sterili conforme. 

Dutch New Guinea : 7 km NE. of Wilhelmina-top, alt. 3,560 m, 
Brass & Myer-Drees 10032, type in Gray Herbarium. "One 
plant in moss of forested cliff." 

Related to E. fuscum. 

11. E. ANGULATUM (Blume) Moore 

E, angulatum (Blume) Moore, Index (1857) 5. 

Lake Habbema region. Brass 1050i, 11036, 110i2, 1H99. 
New to New Guinea. Java; Borneo; Philippines. 

12. E. BRUNNEUM Copel. Plate 10. 

E. brunneum Copel., Univ. Calif. Publ. Bot. 18 (1942) 226. 

Rhizomate repente, 2.5-4 mm crasso, paleis 3-4 mm longis 
lanceolatis attenuatis subintegris squarrosis aterrimis nitidis 
dense vestito; phyllopodiis indistinctis ; stipitibus ca. 1 cm 
remotis, 10-20 cm altis, fusco-stramineis, basibus exceptis glab- 
rescentibus; lamina sterile ca. 17 cm longa, 4 cm lata, apice 
acuta, basi abrupte et breviter cuneata, margine anguste carti- 

406 The Philippine Journal of Science ^^^^ 

laginea, squamulis minutis amorphis fissis sparsa^ superne glab- 
rescente, venis vix conspicuis, lamina fertile ca. 12 cm longa, 
2 cm lata, aliter conforme. 

Dutch New Guinea : 6 km. SW. of Bernhard Camp, Idenburg 
River, alt, 1,200 m, Brass 12808, type in Gray Herbarium; 
epiphyte, near the ground, in rain forest. 

13. E. REPENS Copel. Plate 11. 

E. repens CopeL, Univ. Calif. Publ. Bot. 18 (1942) 226. 

Rhizomate repente, 1.5 mm crasso, paleis atris 2-3 mm longis 
lanceolatis attenuatis integris haud densis vestito; stipitibus 
0.5-2.0 cm remotis, phyllopodiis obscuris ca. 1 cm. longis; 
stipitibus frondium sterilium ca. 4 cm, fertilium 8-14 cm altis, 
stramineis, mox glabratis ; lamina sterile ca. 10 cm longa, 3 cm 
lata, apice rontundata, basi sensim cuneata, breviter decurrente, 
margine angusta deilexo cartilagineo, coriacea, glabrescente, 
venis inconspicuis, lamina fertile minore. 

Dutch New Guinea : 15 km SW. of Bernhard Camp, Idenburg 
River, alt. 1,800 m, Brass 1212i; frequent low epiphyte in mossy 

14. E. FUSCUM Copel. Plate 12. 

E. fuscunt CopeL, Univ. Calif. Cont. Bot. 18 (1942) 226. 

Rhizomate repente, 3-4 mm crasso, paleis fuscis nigrescen- 
tibus 4-5 cm longis ovatis et lanceolatis acuminatis sparse et 
irregulariter ciliatis plerisque appressis dense vestito ; stipitibus 
1-3 cm remotis, phyllopodiis 1-2 cm altis vix distinctis 
dense paleatis, frondium sterilium ca. 10 cm altis, 2 mm crassis, 
sulcatis, stramineis vel fusco-stramineis, paleis sursum decres- 
centibus sparsis, frondium fertilium 15-30 cm altis; lamina 
10-15 cm longa, 2.5-4 cm lata, apice rotundata, basi cuneata 
vix decurrente, coriacea, sordide fusca, margine angustissime 
cartilaginea, squamis atris 1 mm longis et amorphis et trian- 
gularibus grosse ciliato-dentatis sparsa, superne subglabres- 
cente, lamina fertile conforme. 

Dutch New Guinea: Lake Habbema, alt. 3,225 m. Brass 9088, 
type, "plentiful in ground moss of open thickets;" ibidem, epi- 
phytic, Nos. 9S54, 9S55; Bele River, 2,200 m, Nos. 11040, 11071. 

There are several other New Guinea collections of Elapho- 
glossum. In Philip. Jour. Sci. 6C (1911) 92, I reported a 
doubtful E. conforme from Goodenough Bay, King 212; it is not 
that species, lior fit for identification. Better specimens, both 
apparently new, but left undescribed, are King 482, from the 


Copeland: Aspidiaceae of New Guinea 407 

mountains behind Medan; and Brass 4772, from Murray Pass, 
alt. 2,840 m. 

15. DEYOPTERIS Dryander 

Key to the species 

Pinnae subpinnate 1. P. paleacea 

Frond at least tripinnatifid at base. 

Lowest pinnae much enlarged 2. D, sparsa 

Lowest and succeeding pinnae similar. 
Sterile pinnules hardly pinnate. 

Fronds uniform 3. D, bamleriana 

Fronds somewhat dimorphic 4. Z>. pseudoparasitica 

Pinnules freely compound 5. D. suharhorea 

1. D. PALEACEA (Swartz) C. Chr. 

Z>. paleacea (Swartz) C. Chr., Amer. Fern Jour. 1 (1911) 94. 

JBmss 91S8, Lake Habbema, alt. 3,225 m; No. 1028 S, alt. 
2,800 m; No. U09, Mt. Albert Edward, alt 3,680 m. 

At major altitudes in Borneo, Philippines, Formosa, Hawaii, 
India and China, Madagascar, Tropical America. 

2. D. SPARSA (Ham.) O. K. 

D. sparsa (Ham.) 0. K., Rev. Gen. PI. II (1891) 813. 
D. papuana C. Chr., see Brittonia 2 (1987) 298. 

Brass 10248, Bele River, alt. 2,300 m; Brass il28, 4781, 

Papua, Central Division, alt. 2,100 and 1,900 m. The type of 
D. paptuina {Nephrodium dissitifolium Baker) was from Mt. 
Scratchley, alt. 10,000-13,000 feet. 
To India. 


D. bamleriana Ros., Fedde's Repert. 10 (1912) 334. 

The type is Bander 52, Sattelberg, Brass ISO 58, Idenburg 
River, alt. 850 m, is too similar to be distinguished, but is 
apparently without paraphyses. It is an epiphyte, with short- 
creeping rhizome. 


Related to 2). bamleriana, but distinct, is Carr IS 886, received 
from the British Museum (Mr. Alston) as D. Petelotii, 


D. pseudoparasitica v.A.v.R., Nova Guinea 14 (1924) 19. 

Brass 12067, "occasional low epiphyte in mossy forest at 
1,800 m." Frond herbaceous; very similar to D. bamleriana 

408 The Philippine Journal of Science ^^^^ 

except for dimorphism. The type is Lam 19 US, Doorman-top, 
alt. 2,480 m. 

5. D. SVBARBOBEA (Baker) C. Chr. 

P. subarborea (Baker) C. Chr., Index (1905) 295. 

Brause, Engler's Jahrb. 56 (1920) 94, refers to this species 
12 collections by Ledermann. Rosenstock, Fedde's Repert. 12 
(1913) 173, 174, has described varieties: 

attentiataf Keysser S. 18^^ Sattelberg, alt. 800-1,000 m. 
quadripinnata, Keysser B 83 (1912) Bolan, 2,400-3,000 m. 
decornposita, Keysser B 52, Bolan, 2,400-3,000 m. 
biformis, Keysser 39, 40, Bolan, 2,400-3,000 m. 

We have isotypes of vars. qvxidripinnata and biformis; also, 
Bamler, Rosenstock Fil. novog, exsicc. n. 229, labelled simply 
D. subarborea. I do not believe that these are all one species. 
Brass 12107, "a common low-climbing epiphyte in the mossy 
forest at 1,800 m;" seems by description to be var. attemuita. 
The group is a difficult one, of more than one confused species 
in Java (as D. purpurascens) , in Borneo, and in the Philippines, 
as well as in New Guinea; commonly represented in herbaria 
by minor fragments of fronds. 

16. CTENITIS Christensen 

Key to the species 

Frond tripinnatifid or more compound. 

Lowest pair of pinnae much enlarged 1. C. dissecta 

Lower pinnae all similar. 
Stipe sparsely hairy. 

Lamina over 40 cm long 2. C hypolepioides 

Lamina under 15 cm long 3. C alpina 

Stipe densely setose and scaly. 
Sori dorsal on veinlets. 

Frond tripinnate 4. C. vilis 

Frond quinquepinnatifid 5. C spedosissima 

Sori terminal on veinlets 0. C. pulchra 

Frond bipinnate, stipe brown 7. C. habbemensis 

Frond hardly bipinnate, stipe black 8. C. sagenioides 

1. C. DISSECTA (Forster) Copel. 

C. dissecta (Forster) Copel., Genera (1946) 124. 

Clemens 1H22, Morobe, alt. ca. 1,000 m; Bamler R 16, Rook 
Island, alt. 100 m. Reported from Dutch New Guinea by van 
Alderwerelt van Rosenburg, Lam 682, alt. 10 m. I would expect 
it to be common at minor altitudes. 

Polynesia and Malaya. 


Copeland: Aspidiaceae of New Guinea 409 

2. C. HYPOLEPIOIDES (Bos.) Copel. 

C, hypolepioides (Ros.) Copel., Genera (1946) 124. 
Dryopteris hypolipioides Ros., Fedde's Repert. 12 (1913) 175. 

Keysser B 13 (1912), Bolan Mountains, alt. 3,400-3,800 m, 
isotype in Herb. Univ. Calif; Brass 9295, Lake Habbema, alt. 
3,225 m, common floor-fern in subalpine forest. The new collec- 
tion is far more ample than the type, but identical in hairs, 
paleae, residual roughness, and in having vestigial indusia — 
which can be detected on some sori of our isotype. 

The rhizome is ascending, the stipes contiguous but not 
crowded. Stipe 55 cm tall, lamina 80 cm long, lower and medial 
pinnae 25 cm long, 10 cm wide; larger pinnules 6 cm long, 
15 mm wide; secondary pinnules many, free, subpinnate with 
rounded or somewhat cuneate segments, the sterile ones 1.7 mm 
long by 1.3 mm wide, the fertile considerably contracted and 
covered by the sori. Except in stature and consequent dis- 
section, the identity is so complete that I believe this must 
represent in full development the species of which the type 
collection is an ill developed form. Endemic. 

3. C. ALPINA (Eos.) Copel. 

C, alpina (Ros.) Copel., Genera (1946) 124. 
Dryopteris alpina Ros., Fedde*s Repert. 12 (1913) 178. 

Keysser B 36, Bolan, alt. 3,600-3,800 m, isotype in Herb. Univ. 
Calif. This may represent the preceding species, extremely 
reduced at extreme altitude ; if this is the case, the correct name 
is C. alpina. 

4. C. VBLIS (Kunze) Ching 

C. vilis (Kunze) Ching, Bull. Fan Bot. 8 (1938) 290. 

Schlechter 17792, Ibo Mountains, alt. 1,000 m, received as 
Dryopteris intermedia (Blume) 0. K., var. microloba, from 
which (the Mindanao varietal type) it differs only in having 
more acute segments. It is never the original C. vilis than are 
various specimens ascribed to that species in recent years. 

5. C. SPECIOSISSIMA Copel. Plate 13. 

C. speciosissima Copel., Genera (1946) 125. 

Dryopteris speciosissima Copel., Univ. Calif. Publ. Bot. 18 (1942) 219. 

C. gregis C. vilis quinquepinnatifida ; rhizomate erecto, 1-1.5 
cm crasso, apice paleis rufis 1 cm longis 1-2 mm latis integris 
immerso; stipite ca. 75 cm alta, basi nigro-fusco 8 mm crasso, 
sursum ruf o-f usco 4 mm crasso supeme trisulcato, ubique paleis 

410 The Philippine Journal of Science ^^*® 

setiformibus atrocastaneis squarrosis 5 mm longis horrido, et 
paleis aliquot dejectis basibus nigris relictis aspero; lamina 1 m 
longa, deltoideo-ovata, rhachibus paleis setiformibus et setis, 
sursum pilis rubidis nigrescentibus vestitis; pinnis oppositis, 
infimis 35 cm longis, 15 cm latis, acuminatis, pinnula infima 
acroscopica 6 cm longa, sequente basiscopica 11 cm longa, 3.5 
cm lata acuminata bipinnata, pinnulis tertiariis 3 m longis, 
1.5-2 mm latis obtusis inferioribus fere ad costam pinnatisectis, 
segmentis 0.5 mm latis rotundatis herbaceis costis costulisque 
pilis sparsis deciduis ornatis; venis in segmentis simplicibus, in 
pinnulis sequentibus integrioribus vel integris pauci-ramosis ; 
soris ad venulas fere basalibus, indusiis f uscis membranaceis 
mox plicatis, nonnullis orbiculari-reniformibus, aliis ut videtur 

Dutch New Guinea: Bele River, alt. 2,200 m. Brass 11257, 
"in, forest undergrowth, one large clump on a limestone cliff." 

In its deltoid frond and opposite pinnae, this species resembles 
the Peranema-Diacalpe group of genera. Although very thin, 
the indusia persist, but are so roUed-up and distorted that it 
is difficult to be sure of their form. 

6. C. PULCHRA Copel. Plate 14. 

C. pulchra CopeL, Genera (1946) 124. 

Dryopteris pulchra CopeL, Univ. Calif. Pnbl. Bot. 18 (1942) 219. 

Formis dissectis C. vilis similis; rhizomate terrestre, revi- 
repente adscendente, ca. 1 cm crasso, basibusque stipitum paleis 
atrocastaneis lineari-acicularibus integris 1 cm longis vestitis 
et furfuraceis; lamina 45 cm alta, deltoideo-ovata, acuminata, 
basi quadripinnatifida ; pinnis infimis ca. 15 cm longis, basi 
9 cm latis, basiscopice dilatatis; pinnula infima acroscopica, 
sequente proxima basiscopica 5.5 cm longa ad costam alatam 
pinnatisecta segmentis 1 cm longis etiam oblique pinnatisectis 
segmentis secundariis deltoideo-oblongis obtusis vix 1 mm latis ; 
pinnis sequentibus gradatim angustioribus subsessilibus, oppo- 
sitis, pinnulis infimis semper basiscopicis ad alam angustam 
costarum pinnatisectis, segmentis elongato-oblongis plerisque 
integris membranaceis obscure viridibus glabris, costis pinnu- 
larum piluliferis; rhachi frondis conspicue, pinnae inconspicue 
paleaceis ; soris inf ramarginalibus ad venalas apicalibus, indusiis 
persistentibus plerisque orbiculari-reniformibus, sororum supe- 
riorum interdum athyrioideis. 

Dutch New Guinea: 4 km SW. of Bernhard Camp, Idenburg 
River, alt. 850 m, Brass 13i55, type; also. No. 128^8, alt. 1,150 
m, frequent on rocks in rain-forest ravines. 

"^■^'^ Copeland: Aspidiaceae of New Guinea 411 

In spite of some notable differences, especially in the position 
of the sori at the ends of the veinlets, this seems to be related 
to C. vilis, 

7. C. HABBEMENSIS Copel. Plate 15. 

C habbemensis Copel., Genera (1946) 124. 

Dryopteris habbemensis Copel., Univ. Calif. Publ. Bot. 18 (1942) 216. 

C. rhizomate erecto, elongato, 5 mm crasso, nigro, apicem 
versus basibusque stipitum paleis nigro-fuscis ca. 5 mm longis 

1 mm latis acuminatis vestitis ; stipitibus f asciculatis, 25 cm altis, 

2 mm crassis, fuscis, sursum paleis parvis remotis sparsis et 
minute pubescentibus rhachi inconspicue velutina fere epaleata; 
lamina 30 cm alta, 5-9 cm lata, brevi-acuminata, bipinnata; 
pinnis infimis deflexis, 3.5 cm longis, medialibus 4.5 cm longis, 
1 cm latis, acutis basi truncatis sessilibus, rhachillis utraque 
facie pubescentibus ; pinnulis 10-12-paribus, 7 mm longis 2 mm 
latis, obtusis, plerisque adnatis non confluentibus, infimis sessi- 
libus basi lobatis, ceteris obscure crenatis, glabris, firme papy- 
raceis, superne nigro-viridibus inf erne obscure olivaceis ; venulis 
4-5-'paribus, simplicibus, plerisque soriferis; soris medialibus, 
superficialibus, indusiis orbiculari-reniformibus, fuscis, glabris, 

Dutch New Guinea: Lake Habbema, alt. 3,225 m, Brass 9304, 
type, "plentifully scattered over mossy floor of forest; tufts 
±: 60 cm high." No. 9217, same locality, is larger and more lax 
than the type, the lower pinnules relatively long and narrow, 
and more deeply cleft or lobed. No. 11245; Bele River, alt. 
2,300 m, "abundant in openings of ridge forest," is like No. 
9217 except for purplish-black stipes; its suberect caudex is 
25-30 cm tall. The type resembles C. viscosa in size, shape and 
color, but is freely bipinnate. This affinity is more unmistakable 
in the case of No. 11245; but specific identity with C. viscosa 
or any of its described forms or relatives, as I understand them, 
is impossible. 

Brass 11895, Idenburg River region, alt. 1,800 m, has the 
larger pinnae pinnate a third of their length. It is apparently 
an undescribed relative of C. viscosa and C. habbemensis. 

8. C. SAGENIOmES (Mett.) Copel. 

C sagenioides (Mett.) Copel., Genera (1946) 124. 

I mistrust the presence of this species in New Guinea. It 
is listed by Brause, Engler's Jahrb. 56 (1920) 94, as "var. 
Sagenia livida Mett. msc." Bamler 103 (1914), received here 
as Dryopteris sagenioides, is Pteridrys olivacea. 

412 The Philippine Journal of Science ^^^^ 


O. PHAEOSTIGIMA (Cesati) Copel. 

D. phaeoatigma (Cesati) Copel., Genera (1946) 126. 

Aspidium phaeostigma Cesati, Rend. Ac. Napoli 16 (1877) 26, 29. 

Dryopteris phaeostigma C. Chr., Index (1905) 284; Dansk Bot. 

Arkiv 9 No. 3 (1937) 47. 
D. Kingii Copel., Philip. Jour. Sci. 6 C (1911) 73, non C. Chr. (1905) 
D. tamatana C. Chr., Suppl. I (1913) 40. 
Polystichum lastreoides Eos., Fedde's Repert. 9 (1911) 425. 
Dryopteris Ledermanni Brause, Engler's Jahrb. 56 (1920) 90. 
P. eyclosorus v.A.v.R., Nova Guinea 14 (1924) 21. 

Brass 8889, Hollandia, alt. 50 m; 13855, alt. 175 m; U22, 
Aisa River. The most named, and one of the most collected 
New Guinea ferns. 


18. PTERIDRYS Chrlstensen and Ching 

P. OLIVACEA (Ros.) Copel. 

P. oUvaeea (Ros.) Copel., Genera (1946) 126. 
Dryopteris oUvacea Ros., Hedwigia 56 (1915) 352. 

Sattelberg, alt. 800-900 m, Bamler 108 of 1914, received by 
the University of California as Dryopteris sagenioides. It dif- 
fers from P. microthecia (Fee) C. Chr. et Ching by having 
basal pinnae neither enlarged nor dilated, and by having fewer 
sinuses obstructed by teeth. Endemic. 


H. PROFEREOIDES (Christ) Copel. 

H. profereoides (Christ) Copel., Univ. Calif. Publ. Bot. 16 (1929) 61. 
Aspidium profereoides Christ, Philip. Jour. Sci. 2 C (1907) 156. 
A. subasquale Ros., Fedde's Repert. 12 (1913) 176. 
Tectaria subaequalis Copel., Philip. Jour. Sci. 9 C (1914) 5. 

Known in New Guiea only by Bamler's collection or collections 
on the Sattelberg. The type of A. suhaeqimle was collected in 
1912. Our specimen is Rosenstock, Fil. novog. exsicc. n. 2H, 
dated April, 1914, labelled "Aspidium subaequale Ros. n. sp." 
It is less dimorphic than the type collection of H. profereoides, 
but otherwise identical. There is no indusium, and the costal 
sori are oblong. 


S. AURrrA (Swartz) Presl 

S. aurita (Swartz) Presl, Tent. (1836) 237. 

King 285; Brass 5651, Papua. Previously reported. 
Solomon Islands to India. 

'^'^' * Copeland: Aspidiaceae of New Guinea 413 

21. TECTARIA CavaniUes 

This is a difficult genus, for natural reasons, because many 
of its species are ill fixed in the degree of dissection of their 
fronds, and because the indusia are variable and inconstant 
within some of the species. It is artificially difficult in reflection 
of the natural difficulties, because its species have been described 
in a number of unreal genera. Any present treatment of the 
New Guinea species is somewhat tentative because the collec- 
tions are not numerous enough to permit final judgment of 
the local variation within specific limits. 

Key to the species 

Lamina at least bipinnate at base. 
Pinnules at least pinnatifid. 

Areolae along costae and main veins only. 
Pinnules decrescent upward, or few. 
Indusium present. 

Frond green, clear but dark 1. T. devexa 

Frond brownish-green 2. T. Kingii 

Sori naked. 

Lamina subcoriaceous 3. T, andaiensis 

Lamina thin. 

Rachises scaly 4, T. ferruginea 

Rachises puberulous 5. T. gymnocarpa 

Pinnules numerous and uniform 6. T, leuzeana 

Veins anastomosing freely. 

Sori naked 7. T. irregularis 

Indusium present. 

Lamina coriaceous, naked 8. T. Ledermanni 

Lamina thin, pubescent ,. 9. T. pubescens 

Pinnules not pinnatifid. 
Indusium persistent. 

Fertile suprabasal pinnae pinnatifid 10. T, cesatiana 

Fertile suprabasal pinnae crenate 11. T. Barclayi 

Indusium fugacious 18. T. melanocaulis 

Lamina simple to pinnate, not minute. 
Stipe not winged. 
Pinnae normally more than 3 pairs. 
Fronds clustered. 
Sori round. 
Pinnae broad at base. 

Indusium persistent 11. T. Barclayi 

Indusium fugacious 19. T. Weinlandii 

Pinnae narrow at base. 

Indusium reniform 12. T, erenata 

Indusium peltate 13. T, tematensis 

Sori suboblong 14. T. pleiosora 

Sori amorphous, exindusiate 15. T, teratoearpa 

414 The Philippine Journal of Science ^^^9 

Bhizome elongate, fronds ieriate. 

Pinnae 3 cm wide, not forked 16. T. Menyanthidis 

Pinnae 2 cm wide, the lower forked 17. T. aemibipinnata 

(Frond simple or with 1 to 8 pinnae) 
Frond simple or with 1 to 2 pinnae. 

Pinnae pinnatifld 20. T. pleocnemioides 

Margin broadly crenate 21. T. novoguineensis 

Margin essentially entire. 

Indusium none or inconspicuous. 
SoTi scattered. 

Sori minute, somewhat irregular 22. T. angulata 

Sori irregularly elongate 23. T. cristovalensis 

Sori in paired rows 24. T, siifolia 

Indusium conspicuous. 

Sori along main veins 12. T. crenata 

Sori submarginal 25. T. eraspedocarpa 

Rachis and stipe broadly winged. 

Sori in paired rows 28. T. decurrens 

Sori scattered 26. T. heccariana 

Lamina simple, under 4 cm long 27. T. minuta 

1. T. DEVEXA (Kunze) Copel. 

T. devexa (Kunze) Copel., Philip. Jour. Sci 2 C (1907) 415. 

Accredited to New Guinea as Pleocnemia membranxicea Bedd., 
var. novoguineensis Eos., Fedde's Repert. 10 (1912) 338, listed 
by Brause, Engler's Jahrb. 56 (1920) 116, Aspidium devexum, 
var. novoguineensis Ros. We have this plant, Rosenstock Fil 
novog. exsicc. n. 68. It is positively not Aspidium membmna- 
ceum Hooker. It is near T. gigantea (Blume) Copel., but hardly 
identical. It is larger than T. Kingii, with broader segments, 
and is dark-green, not brownish-green. I list T. devexa because 
this plant is not to be listed elsewhere, and I do not wish to 
describe our specimen as new. 

Aspidium devexum Kunze was a nomen nudum, a substitute 
for ''Sagenia intermedia J. Sm. (Cuming. Phil No. 177), ''—also 
a nomen nudum, the more mysterious because 177 was not a 
Cuming collection number, the plant having none. I construe it 
by Aspidium membranaceum Hooker, Sp. Fil. V 105. 

2. 7. KINGn Copel. 

T. Kingii Copel., Philip. Jour. Sci. 9 C (1914) 4. 

Known only by the original collection, King i02, from Wood- 
lark Island. 

3. T. ANDAIENSIS (Baker) C. Chr. 

T. andai$nsis (Baker) C. Chr., Suppl. Ill (1934) 177; see Dansk 
Bot. Arkiv 9 No. 3 (1937) 49. 

'^'^'^ Copeland: Aspidiaceae of New Guinea 415 

Known only by the original collection, Herb. Beccari No, 

4. T. FERRUGINEA (Mett.) Copel. 

T. f&rmginea (Mett.) Copel., Philip. Jour. Sci. 6 C (1911) 76. 

Brass 88S2, HoUandia, alt. 50 m. 

Described from a collection by Zippelius. King 264, from 
Papua, fits Mettenius' description, but differs from his figure 
in being somewhat more dissected. 


5. T. GYMNOCARPA Copel. 

T. gymnocarpa Copel., Philip. Jour. Sci. 9 C (1914) 4. 

Known only by the original collection, King iOl, from Loana, 
Papua. Christensen reduces this to T. ferruginea, calling my 
attention to Mettenius' figure. There are resemblance and af- 
finity; but King iOl and 264 differ essentially in paleae and 
hairs, carefully described in Mettenius' text, but omitted on 
his illustration. 

6. T. LEUZEANA (Gaud.) Copel. 

jT. leuzeana (Gaud.) Copel., Philip. Jour. Sci. 2 C (1907) 417. 

Brass 129S8, alt. 1,200 m; 1S857, alt. 130 m. 

Common but variable. Rosenstock has described varieties 
echinocarpa and lobato-crenata; and a Winkler collection received 
from Rosenstock as Aspidium chrysotrichum Baker, var, can be 
included here. However, more than one distinct species may 
already be included. 

Polynesia to Asia. 

7. T. IRREGULARIS (Presl) Copel. 

r. irregularis (Presl) Copel., Philip. Jour. Sci. 2 C (1907) 416. 
Common. Fiji to Asia. 

8. T. LEDERMANNI (Brause) C. Chr. 

T, Ledennanni (Brause) C. Chr., Suppl. Ill (1934) 181. 

Not seen; known only by the original collection, Ledennann 
9409, from the Sepik region, alt. 850 m. 

9. T. PUBESCENS Copel. Plate 16. 

T. pubescens Copel., Univ. Calif. Publ. Bot. 18 (1942) 221. 

Rhizomate et parte inferiore stipitis ignotis; stipite 4 nmi 
crasso, facie anteriore trisulcato, brunneo-pupurascente, rhachi- 
busque pilis minutis articulatis debilibus sat dense pubescentibus ; 

10882 4 

416 The Philippine Journal of Science ^^^^ 

lamina ultra 50 cm longa, deltoideo-ovata, basi imo tripinnata; 
pinnis infimis fere 30 cm longis, pinna infima basiscopica 12 cm 
longa, ad alam apicem versus latam deorsum contractam pinna- 
tifida, pinnula secundaria libera una, segmentis sequentibus pin- 
natilide lobatis 13 mm latis, lobis 3-4 mm latis obtusis; pinnis 
sequentibus 20 cm longis, vix obliquis, deorsum leviter bipinna- 
tifidis; pinnis superioribus pinnatifide lobatis, deinde in apicem 
sat magnum coadunatis; lamina herbacea, utraque facie pube- 
scente; venis laxe anastomosantibus, utroque latere costae seg- 
menti seriem unam areolarum majorum et hie illuc seriem alte- 
ram minorum includentibus ; soris parvis irregulariter seriatis, 
medialibus, aut ad apices venularum inclusarum aut apices versus 
loborum omnino liberarum, indusio typi persistente, plerumque 
cordato rarius peltato, glabro. 

King 359, Lakekamu, Papua, type, — reported, Philip. Jour. 
Sci. 6 C (1911) 76, as T. malayensis; also, Bamler, Rosenstock 
Fit novog. exaicc. n. 236, received as Aspidium coadunatum 
Wall. var. nifovillosa Ros. n. v,, from Logaueng, Kaiser-Wil- 

A relative of T. ferruginea, less dissected, and therefore with 
more areolae, and indusiate. The Bamler plant is old, and but 
few indusia can be detected. On the type, but not on the 
Bamler plant, there are a few narrowly linear paleae 1-5 mm 
long on the rachises. 

As the specimens cited as T. pubescens are the only bases for 
reports of T. malayensis and T. coadunata in New Guinea, these 
two species are dropped from the list of of New Guinea ferns. 

10. T. CESATIANA (C. Chr.) Copel. 

T. cesatiana (C. Chr.) Copel., Philip. Jour. Sci. 6 C (1911) 76. 
Aspidium bamlerianum Eos., Fedde's Repert. 10 (1912) 330. 
Tectaria bamleriana C. Chr., Suppl. Ill (1934) 177. 

Brass 13408, alt. 850 m, largest frond 13 cm long, subpinnate ; 
No. 13H2, alt. 800 m, largest frond 43 cm long, tripinnatifid at 
base; No. 13852, alt. 170 m, up to 50 cm long. More or less 
dimorphic, usually conspicuously so. 

Our specimen of A. bamlerianum, received from Rosenstock, 
and bearing his notation "unicum," is exactly this species. 

Throughout New Guinea. Endemic. 

11. T. BABCLAYI (Carr.) C. Chr. 

T. Barclayi (Carr.) C. Chr., Suppl. Ill (1934) 177. 

Carr llOU, 11U5, 12334, det. et misit Alston, Brass 3648 
and 5572, from Papua, det. Christensen as T. papuuna Copel., 

'^'^'^ Copeland: Aspidiaceae of New Guinea 417 

are this species. The type of T. papwana looks distinct, the 
margin being entire to crenate, the intermediate pinnae broadly 
rounded at base and not at all pronged; but, so variable in 
dissection are the species of this genus, it may be only an 
undissected form, T. Barclayi shows affinity to T. crenata, 
which the type of T. papuana does not suggest. 
New Ireland. 

12. T. CRENATA Cav. 

r. crenata Cav., Descr. (1802) 250. 

Brass 12222, alt. 1,800 m; No. 18834, deltoid and trifid. In 
most genera, the latter specimen would be accepted without 
question as specifically distinct, but even great differences in 
form and dissection are not by themselves safe criteria in 
Tectaria, Conspecific with this specimen is Bamler H of 1913, 
received from Rosenstock as Aspidium grandifolium Presl, which 
it is not; it has free, unforked basal pinnae and one pair of 
almost free lateral pinnae. These specimens may represent a 
distinct species, but are at any rate near T. crenata. 

Common. Asia to Polynesia. 

13. T. TERNATENSIS v.A.v.R. 

T. ternatensis v.A.v.R., Bull. Dept. Agric. Ind. Neerl. No. 18 (1908) 9. 

Reported by van Alderwerelt, Nova Guinea 14 (1924) 8. 

Distinguished by its author from Aspidium repandum Willd. 
by size and by narrower pinnae; from A. persoriferum Copel. 
and A. pachyphyllum Kunze by the shape of the indusium. 
These are all regarded as forms of T, crenata, and any real 
distinction of A. or T. ternatensis is not evident. 

14. T. PLEIOSORA (V.A.V.R.) C. Chr. 

T. pkiosora (v.A.v.R.) C. Chr., Gardens' Bull 7 (1984) 260. 

To be distinguished from T. crenata by having at least a 
part of the sori oblong instead of round, and fewer, broader 
pinnae, broader particularly on the basiscopic side. Regarding 
T. crenata as a freely variable species, I mistrust the distinc- 
tions. I have seen no New Guinea specimen, but have an 
authentic one from Borneo. 

15. T. TERATOCARPA (v.A.v.R.) C. Chr. 

T. teratocarpa (v.A.v.K.) C. Chr., Suppl. Ill (1934) 185. 

Known only by the original collection. Lam 966, from the 
Mamberamo River, alt. 100 m; isotype in Herb. Univ. Cali- 

418 The Philippine Journal of Science ^^^^ 

16. 7. MENYANTHIDIS (Presl) Copel. 

T. Menyanthidis (Presl) Copel., Philip. Jour. Sci. 2 C (1907) 414. 

Brass 13860, alt. 150 m; King 169, 180; Brass 975; apparently 
Philippines; Admiralty and Solomon Islands. 

17. T. SEMmiPINNATA (Wall.) Co]»eI. 

T. semibipinnata (Wall.) Copel., Sarawak Mus. Jour. 2 (1917) 371. 

What is reported from New Guinea is Aspidium nudum 
(Baker) Diels, said to be a synonjntn of T. semibipinnata, its 
type from Borneo. The resemblance to T. Menyanthidis is so 
close that confusion is possible. 

18. T. MELANOCAUUS (Blume) Copel. 

jT. melanocaulis (Blume) Copel., Philip. Jour. Sci. 2 C (1907) 416. 

Our only New Guinea specimen is Bamler, Rosenstock FiL 
novog. exsicc, n. 117, from Wareo, alt. 600 m. 
Malaya; Philippines; reported from China. 

19. T. WEINLANDII (Christ) Copel. 

T. Weinlandii (Christ) Copel., Jour. Arnold Arb. 10 (1929) 177. 

Central and Eastern New Guinea, at minor altitudes. 

20. T. PLEOCNEMIOIDES (v.A.v.R.) C. Chr. 

T. pleocnemioides (v.A.v.R.) C. Chr., Suppl. Ill (1934) 183. 

Known only by the type collection. Lam i2S, Mamberamo 
River, alt. 25 m. 

21. T. NOVOGUINEENSIS (Ros.) C. Clir. 

T. novoguineemis (Ros.) C. Chr., Suppl. Ill (1934) 182. 

Known only by an old collection by Zippelius in the Rijks 
Herbarium, Leiden, very imperfectly described: "Ab Aspidium 
Lahrusca Christ stipitibus pilosis, laminis grosse crenatis, in- 
dusio magno, persistente diversum.'' 

22. T. ANGULATA (WlUd.) Copel. 

T. angulata (Willd.) Copel., Sarawak Mus. Jour. 2 (1917) 370. 

Brass 8886, HoUandia, alt. 50 m. Previously collected by 
Schlechter and Bamler. 
Malaya ; Philippines ; Solomon Islands. 

23. T. CRISTOVAOn^SIS (C. Chr.) Alston 

T. cnstovalensis (C. Chr.) Alston, Jour. Bot. (1939) 290. 

Brass IS 85 8, Idenburg River, alt. 80 m. Reported from New 
Guinea as Dictyopteris pentaphylla v.A.v.R., Aspidium quinque- 

"^"^'^ Copeland: Aspidiaceae of New Guinea 419 

foliolatum C. Chr., Campylogramma pteridiformis v.A.v.R., and 
Tectaria diversisora Copel. 
Solomon Islands. First named Gymnogramme palmata Baker. 

24. T. SmJ'OLIA (Willd.) Copcl. 

r. siifolia (Willd.) CopeL, Philip. Jour. Sci. 2 C (1907) 414. 

Reported from New Guinea, but I have not seen it. 
Malaya; Philippines. 


r. craspedosora CopeL, Jour. Arnold Arb. 10 (1929) 178. 

Known only by the type collection. Brass 557, Laloki River, 
alt. 1,500 feet. 

26. T. BECCARIANA (Cesati) C. Chr. 

T. beccariana (Cesati) C. Chr., Suppl. Ill (1934) 177; see Dansk 

Bot. Arkiv. 9 No. 3 (1937) 51. 
Polypodium ingens Brause, teste Christensen, 1. c. 
Aspidium vaatum Blume forma latins v.A.v.R., Nova Guinea 14 

(1924) 7. r. vasta (Blume) Copel., is otherwise unreported from 

New Guinea. 

Mindanao, as IT. Bryanti Copel. 

27. T. MINUTA Copel. 

T. minuta Copel., Philip. Jour. Sci. 30 (1926) 328. 
Known only by the type collection. King i93, 1. Rev. P. C. 
Shaw, Mountains behind Taupota. 

28. T. DECURRENS (Presl) Copel. 

T. decurrens (Presl) Copel., Elmer's Leaflets Philip. Bot. 1 (1907) 234. 

Brass 12899, IS 85 4, IS 85 5 A, 1S85S. The last, a single plant, 
has simple fronds, entire or broadly and irregularly crenate, 
attenuate to a short, scaly stipe. Found with typical plants of 
the species, Nos. 1385^ and 13855 A, it may be regarded with 
confidence as an abnormal, or probably juvenile, individual. Our 
specimen of Lam 1323, Doorman River, alt. 200 m, reported. 
Nova Guinea 14 (1924) 13, as approximately Dictyopteris 
heterosora (Baker) Bedd., is T. decurrens. 

Common. Polynesia to India. 

22. HEMIGRAMMA Christ 
H. HOLLRUNGU (Kuhn) Copel. 

H. Hollrungii (Kuhn) Copel., Philip. Jour. Sci. 37 (1928) 406. 

Besides the original collection, Brause cites Ledermann 6562, 
7781. This species was described as having pinnatifid fronds. 

420 The Philippine Journal of Science ^^^^ 

To it, Christensen, SuppL III 109, reduces H. grandifolia Copel, 
with pinnate sterile frond, the rachis mostly wingless. This 
may be correct, as Hemigramma is a recent derivative of 
Tectaria with pinnate fronds, and reversions by species with 
condensed, even with simple and entire fronds, are known to 
occur. H. grandifolia looks like the most primitive element in 
its genus. The range of the genus is to Java, China and 
Formosa; and it is in harmony with my theory as to the 
direction of migration in this region that the most primitive 
member of a genus with this range should be found in New 
Guinea. The type of H, grandifolia is King 828, from Lake- 
kamu, Papua. 
New Britain. 

23. LASTREA Boiy 

Key to the species 

Frond tripinnatifid at base or more compound. 
Eachises and costae not scaly. 

Stipe smooth 58. L. setosa 

Stipe muricate 59. L. armata 

Rachises bearing scales and hairs 60. L, leucolepis 

Frond bipinnate. 
Pinnae freely pinnate. 

Frond not over 60 cm long. 

Sori submarginal 2. L. bipinnata 

Sori costular 7. L, conterminoides 

Frond several meters long 1. L. marattioides 

Free pinnules only about 3 pairs 3. L. platyptera 

Pinnae pinnate at base only or not at all. 

Veinlets typically forked 4. L, Jiavo-virens 

Veinlets simple. 
Lower pinnae gradually dwarfed. 
Largest pinnae under 5 cm long. 
Rachis sparsely hairy or naked. 

Rhizome creeping, slender 5. L. Beddomei 

Stem erect, stouter 6. L, Brassii 

Rachis persistently pubescent. 
Pinnae deeply pinnatifid. 

Sori subcostular 7. L. conterminoides 

Sori medial 8. L. pefpahescens 

Pinnae shallowly pinnatifid 9. L. petrophila 

Pinnae 10 cm or more long 10. L. Harveyi 

Lower pinnae abruptly reduced. 
Lower pinnae over 20 cm long. 

Pinnae cut to 1 mm from costa 11. L. subattenuata 

Pinnae pectinate to a broad wing 12. L, pseudostenobasis 

Pinnae about 15 cm long 13. £,. Regis 

Larger pinnae 7-10 cm long. 

■^^'^ Copeland: Aspidiaceae of New Guinea 421 

Chartaceous, glabrous beneath 14. L. novoguineensis 

Herbaceous, glandular beneath 15. Dryopteris glauces- 

Pinnae under 5 cm long. 

Rachis minutely pubescent 16. L. Finisterrae 

Rachis hairy and scaly 17. L, mixta 

Lower pinnae not or but little reduced. 
Stipe reddish or purplish-black. 
Rachis hairy and paleate. 

Lamina pallid, indusia purplish 18. L. pallescens 

Lamina dark, indusia brown. 

Sori about medial 19. L. viscosa 

Sori nearer the costule. 

Papyraceous 20. L. engleriana 

Coriaceous 21. Dryopteris villosipes 

Rachis hairy, not scaly. 

Membranaceous, diaphanous 22. L. diaphana 

Thin-chartaceous or herbaceous. 

Indusium naked 23. L, subnigra 

Indusium glandular-puberulent 24. Dryopteris oligole- 


Coriaceous 25. L, coriacea 

Stipe stramineous to brown, 

Sori exindusiate, elongate 26. Dryopteris stellato- 

Sori exindusiate, round. 
Sori about medial. 

Veinlets not connivent 27. L. wariensis 

Lowest veinlets connivent 47. L, subdimorpha 

Sori nearer to costule. 
Sporangia naked. 

Stipe not scaly 28. L. tuberculata 

Stipe scaly at base. 

Pinnae up to 5 cm wide 29. L. notabilis 

Pinnae under 3 cm wide. 

Paleae nigrescent 30. L. dryopteroidea 

Basal paleae brown 31. L. varievestita 

Sporangia setose. 

Lamina 70 cm long 32. L. alta 

Lamina up to 35 cm long. 

Pinnae broad-lanceolate 33. L. quadriaurita 

Pinnae narrow-lanceolate 34. L. ensipinna 

Sori indusiate. 
Sori supra-medial to margina}. 

Indusium not hairy 36. L. immersa 

Indusium hairy. 

Hairs of rachis various 35. Dryopteris quadri- 

Hairs of rachis uniform. 

Pinnae 4 cm wide 37. L. keysseriana 

(Pinnae up to 2 cm wide) 
Pinnae up to 2 cm wide. 

422 The Philippine Journal of Science 1849 

Basal pinnae horizontal 38. L, obliquata 

Basal pinnae deflexed 39. Dryopteris Peekeli 

Sori medial. 
Frond thin. 

Stipe scaly at base only 40. L. gracilescens 

Stipe siibscaly upward 41. L. Lauterbachii 

Frond coriaceous. 

Segments 1-2 mm wide 42. Dryopteris stereo- 

Segments 8 mm wide. 

Bachis naked beneath 43. Dryopteris rigidifo- 


Bachis hairy 44. Dryopteris horizon- 

Sori inframedial to costular. 
Lamina thin. 

Bachis hairy 45. L. calcarata 

Bachis naked beneath 46. L. fulgens 

Lamina firm to coriaceous. 
Indusium setulose. 

Bachis pubescent only 48. L. falcatipinnula 

Bachis hairy and scaly. 

Pinnae 15 mm wide 49. L. echinata 

Pinnae 3 cm wide 50. L. helensis 

Indusium naked. 
Pectinate pinnae many. 

Segments 5 mm wide 61. L. costulisora 

Segments 3 mm wide 52. L, ehlamydophora 

Pinnae up to 10 pairs. 
Bachis pubescent. 

Pinnae coriaceous 53. L. ophiura 

Pinnae subcoriaceous 54. L. crassa 

Bachis naked beneath 55. L. wantotensis 

Pinnae nearly or quite entire 50. L, nephrolepioides 

Frond pinnatifid except for basal 
pmnae 57, £,, hunsteiniona 

1. L. MARATTIOIDES (Alston) Copel. 

Dryopteris marattioides Alston, Jour, of Bot. 78 (1940) 227; Nova 
Guinea N. S. 4 (1940) 110, PL 7, /, 7, 8. 

Clemens 4809, type, Morobe, alt. 5,800 feet; 6m8, Sambanga, 
alt. 6,000 feet. Twelve to 14 feet tall. Mrs. Clemens made yet 
a third collection of this remarkable fern, which may well 
represent an undescribed genus. 


2. L. BIPINNATA Copel., comb. nov. 

Dryopteris bipinnata Copel., Philip. Jour. Sci. 9 C (1914) 2. 

Known only by the type collection, King 407, from Loane, 

'^'^''^ Copeland: Aspidiaceae of New Guinea 423 

3. L. PLATYPTEEA Copel. Plate 17. 

L. platyptera Copel., Genera (1946) 139. 

Dryopteris platypUra Copel., Univ. Calif, Publ. Bot. 18 (1942) 219. 

Rhizomate erecto, basibusque confertis stipitum paleis ovatis 
acutis integris glabris fuscis nigrescentibus ca. 1 mm longis 
vestitis; stipite alibi glabro atropurpureo nitente; lamina 30 cm 
longa, 11 cm lata, sensim acuminata, basi vix angustata, rhachi 
atropurpurea costisque superne purpureostrigosis, alibi lamina- 
que glabris; pinnis ca. 13-paribus, inferioribus breviter (1 mm) 
pedicellatis, infimis pauUo abbreviatis vix deflexis, sequentibus 
ca. 5.5 cm longis 2 cm latis, acuminatis, infra apicem integrum 
gradatim profundius pinnatifidis, rhachin versus pinnatis; pin- 
nulis ca. 3-paribus, adnatis vel infimis sessilibus, infima acrosco- 
pica inciso-dentata, sequentibus obscure serratis ca. 11 mm 
longis, 2-3 mm latis, acutis vel obtusis, herbaceis; venis ca. 
6-paribus, infimis saepe f urcatis ceteris simplicibus ; soris inf ra- 
medialibus, parvis, indusiis tenuibus, nudis, deciduis. 

Dutch New Guinea: Bele River, alt. 2,200 m. Brass 11328; 
"in semi-shade on high rocky bank of stream." 

Apparently related to L. pyrrhorkachis, and with resemblance 
but no probable affinity to Ctenitis sagenioides and Lastrea 

4. L. FLAVO-VIRENS (Ros.) Copel. 

L. flavO'Virens (Ros.) Copel., Genera (1946) 138. 
Dryopteris flavo-virens Ros., Fedde's Repert. 10 (1912) 334. 

Bamler, Rosenstock Fil. novog. exsicc, n. 118, Wareo, alt. 
600 m; isotype in Herb. Univ. Calif. Peculiar in Lastrea 
because of many forked veinlets. 

5. L. BEDDOMEI (Baker) Bedd. 

L. Beddomei (Baker) Bedd., Ferns Brit. India Corr. II (1870). 

Brass 9319, 10888, 10932, alt. 2,620-3,225 m; reported by 
Gepp from the Arfak Mountains, alt. 7,000 feet. 

The Brass specimens are identical with those from Java, 
Luzon, Formosa and Yunnan. Stipe and rachis rather decidu- 
ously hairy, hairs more persistent on minor axes; sori submar- 
ginal, indusia glandular or glandular-marginate, transient. The 
typical plant may be represented by Beddome, Ferns of Southern 
India, Plate CX, figured with a more hairy rachis, and described 
as with glabrous indusium. Baker, in publishing the specific 
name, Syn. Fil. (1873) 267, cited Thwaites C. P. 1287, but did 
not mention the rachis or indusium. 

424 The Philippine Journal of Science ^^^^ 

6. L. BRASSII Copel., comb. nov. 

Dryopteris Brassii C. Chr., Brittonia 2 (1937) 295. 

Brass A9S7, Mt. Tafa, alt. 2,400 m. A young frond of the 
isotype in Herb. N. Y. Bot. Garden bears glandular-marginate 
indusia, wanting on older fronds. It still seems distinguishable 
from the preceding species by its erect, scaly rhizome. The 
creeping rhizome of L. Beddomei is characteristic. But L. 
Brassii grew on the "banks of a small stream flowing over debris 
of a landslip." If an established plant grew up through any 
kind of a subsequent deposit, its rhizome would almost have to 
become erect, for the occasion. 

7. L. CONTEBMINOIDES €opel.» comb. nov. 

Dryopteris conterminoides C. Chr., Index (1905) 258. 
Nephrodium simulans Baker, Jour. Bot. 28 (1890) 106, non (1874) 
nee (1888). 

Known only by the original collection by MacGregor, on Mt. 
Knutsford. Described as with densely crinite rachis, subcostal 
sori, and persistent indusium. 

8. h. PERPUBESCENS (Alston) Copel., comb. nov. 

Dryopteris perpubescens Alston, Jour. Bot. 78 (1940) 227; Nova 
Guinea N. S. 4 (1940) 111, PL 8, f. 9, 10, 

Clemens 7902c, Morobe District, Kalasa, alt. 1,600 feet. Not 
seen. Stipe densely puberulous, lamina densely short-pubescent, 
sori medial, indusia hirsute. 

9. L. PETROPHBLA Copel. Plate 18. 

L. petrophila Copel., Genera (1946) 139, 

Dryopteris petrophila Copel., Univ. Calif. Publ. Bot. 18 (1942) 220. 

L. rhizomate suberecto, f rondibusque, circinnatis paleis f uscis 
vix 1 mm longis protectis; stipitibus laxe fasciculatis, 5-7 cm 
altis, gracilibus, brunnescentibus, basi paleis paucis cito caducis 
fere 2 mm longis ovatis sparsis, mox omnino glabris; lamina 
40 cm longa, parte tertia superiore longe attenuata pinnatifida 
vel apicem versus tantum inciso-serrata, parte tertia inf eriore 
etiam arigustata pinnis decrescentibus remotis inflmis minutis, 
parte mediale usque ad 7.5 cm lata subbipinnatisecta, rhachi 
minute pallide velutino-pubescente; pinnis suboppositis, sessi- 
libus, maximis 4 cm longis 8 mm latis, acutis, basi acroscopice 
inconspicue auctis, plus minus medio ad costam inciso-lobatis, 
superne saturate viridibus, costis costulisque minutissime setu- 
losis, inferne pallidis fere (oculo nudo, omnino) glabris; venis 

''^^ * Copeland: Aspidiaceae of New Guinea 425 

simplicibus, infimis ad sinus conniventibus rarius confluentibus ; 
soris medialibus infimis inf ramedialibus, plerisque nudis, rarius 
indusiis vestigialibus f ugacibus praeditis. 

Dutch New Gunea: Bele River, alt. 2,200 m, Brass 11826, 
''abundant on moist rock faces along a forest stream; fronds 

With L. Brassii, this species belongs in a group more diversely 
developed in the Philippines than, so far as now known, in 
New Guinea. From its neighbors in this enumeration, it is 
distinguishable by the shallowly cut pinnae, with consequent 
occasional anastomosis of the lowest veinlets. L. nephrolepio- 
ides, with practically entire pinnae, is also a relative. 

10. L. HARVEYI Carr. 

L. Harveyi Carr., Flora Vitiensis (1873) 359. 

Listed from New Guinea by Brause, but not known to me here. 
Described from Fiji; reported elsewhere in Poljmesia, and 
from the New Hebrides and Admiralty Islands. 

11. L, SUBATTENUATA (Ros.) Copel. 

L. subattenuata (Eos.) Copel., Genera (1946) 1940. 
Dryopteris subattenuata Ros., Fedde's Kepert. 10 (1912) 332. 

Known only by the original collection, Bamler L 37, Rosenstock 
Fil. novog. exsicc. n. 80, from Logaueng, alt. 300 m; isotype 
in Herb. Univ. Calif. 


Dryopteris pseudostenabaais Copel., Jour, Arnold Arb. 10 (1929) 176. 

Brass 1000, Ihu, Vailala River; known by the single collection ; 
possibly a form of Cyclosorus alatellm. 

13. L. BEGIS Copel. Plate 19. 

L. Regis Copel., Genera (1946) 140. 

Dryopteris Regis Copel., Univ. Calif. Publ. Bot. 18 (1942) 220. 

L. rhizomate et stipite ignotis; frondis parte visa 120 cm 
longa, deorwum abrupte contracta, pinnis hie 4-4.5 cm inter 
se distantibus 6-paribus visis, 1-1.5 cm longis, orbiculari-ovatis 
rotundatis, lobatis vel pinnatifidis segmentis ca. biparibus 4 mm 
latis, rhachi 5 mm crassa fusco-plumbeo nudo; pinnis sequentibus 
permultis oppositis, 3-4 cm remotis, sessilibus et tuberculis 
subtensis, ca. 15 cm longis, 2.5 cm latis, acuminatis, basi pauUo 
dilatatis, ad alam 1 mm latam costae suboblique pectinatis, 
costis superne velutinis inf erne glabris; segmentis ca. IS mm 
longis 4 mm latis, rotundatis, integris, papyraceis, glabris; 

426 The Philippine Journal of Science ^^^^ 

venis ca. 8-paribus, simplicibus, infimis ad marginem supra 
sinum currentibus ; soris multis, parvis, subcostularibus, indusiis 
invisis, sporangiis in vetustat enudis. 

Papua: Mountains behind Medan, Rev. Copland King 486. 

The specimen consists of 120 cm of the upper part of a frond, 
wanting the base. The absence of an indusium may be due to 


Dryopteris munda Bos. Meded. Kijks Herb. No. 31 (1917) 5. 

"Nova Guinea: Atasrip 1903." Not seen. So far as the 
very incomplete description shows, this may resemble L. Regis 
except for having hairy axes. 

14. L. NOVOGUINEENSIS (Braiise) Copel. 

L, novoguineensis (Brause) CopeL, Genera (1946) 139. 
Dryopteris novoguineensis Brause, Engler's Jahrb. 49 (1912) 26. 

Brass 12849, alt. 1,200 m. The type is Schlechter 17719, 
Kani Mountains, alt. 100 m. Bamler, Rosenstock Fil. novog. 
exsicc. n. 227, distributed as this species, fits better the descrip- 
tion of Dryopteris glaucescens. 



Dryopteris glaucescens Brause, Engler's Jahrb. 56 (1920) 85. 

Sepik region, alt. 1,400-1,500 m, Lederntann 13034. 
This and the preceding species seem to be very similar. 

16. L. FIHISTERRAE (Brause) Copel. 

L, Finisterrae (Brause) CopeL, Genera (1946) 138. 
Dryopteris Finisterrae Brause, Engler's Jahrb. 49 (1912) 20. 

Schlechter 18134, Finisterre Mountains, alt, 1,000 m; Bamler 
116 (1914), Sattelberg. 

17. Ii. MIXTA (Ros.) Copel., comb. nov. 

Dryopteris mixta Ros., Fedde's Eepert. 12 (1913) 172. 

Known only by the type collection, Keysser S 141, Sattelberg, 
alt. 800-1,000 m; not seen. 

18. t>. PALLESCENS (Brause) Copel., comb. nov. 

Dryopteris paUeseens Brause, Engler's Jahrb. 56 (1920) 88. 

Known only by the type, Ledermann 9146, Sepik region, alt. 
850 m; not seen. Two meters tall. 

19. L. VISCOSA J. Smith 

L. viseosa J. Smith, Jour. Bot. 3 (1841) 412 nomen. 
Nephrodium viscosum Baker, Syn. Fil. (1867) 264. 

^^•^ Copeland: Aspidiaceae of New Guinea 427 

Brass 132H, alt. 900 m, common ground fern in Agathis 
forest. Except that few glands can be detected, this is perfectly 

Philippines; Borneo; and (not quite typical) Fiji. 

This is a variable species or a closely related group. Christen- 
sen has reduced to it two which seemed distinct species to me ; 
and they are indeed reducible if one construe it broadly. If this 
policy be adopted, I mistrust the specific identity of all of the 
following six "species." 

20. L. ENGLEBIANA (Brause) Copel. 

L. engleriana (Brause) Copel., Genera (1946) 138. 
Dryopteris engleriana Brause, Engler's Jahrb. 49 (1912) 19. 

The type, not seen, is Schultze 330, from the Sepik region. 
We have received with this name Keysser 181, Sattelberg 
Hinterland; also, as var. Mrta C. Chr., Brittonia 2 (1937) 296, 
Brass 5032 and UO^. Brass 10281, Lake Habbema, alt. 2,800 
matches these well. All have persistent indusia; the species 
was described as exindusiate. They are also alike in having 
nigrescent stipe-bases, not provided in the description. 


Dryopteris villosipes Gepp. in Gibbs, Dutch N. W. New Guinea 
(1917) 70. 

Gibbs 5627, Arfak Mountains, alt. 7,000-8,000 feet; not seen. 
Epiphytic. Without other comparison, Gepp remarks: "The 
plant differs in every respect from D. viscosa. 

22. L. OIAPHANA (Brau$e) Copel.^ comb. nov. 

Dryopteris diaphana Brause, Engler's Jahrb. 56 (1920) 80. 
Ledermann 8903, Sepik region, alt. 850 m ; collected but once. 

23. L. SUBNIGRA (Brause) Copel. 

L. suhnigra (Brause) Copel., Genera (1946) 140. 
Dryopteris suhnigra Brause, Engler's Jahrb. 56 (1920) 82. 

Ledermann 11962, type, not seen, Sepik region, alt. 2,170 m, 
epiphytic in masses of moss; Clemens il020, Morobe, alt. 1,800 
m. Brass 12038, alt. 1,750 m, a single specimen, epiphytic 
with pendent fronds, is like L. subnigra in most respects, but 
the nether surface is minutely pubescent. 


Dryopteris oligolepia v.A.v.R., Nova Guinea 14 (1924) 17. 

Typified by Lam 1977, Doorman4op, alt. 2,520 m, epiphytic; 
not seen. Because of relative nakedness of stipe and rachis, 
Brass 9062, Lake Habbema, alt. 3,225 m, may be identified as 
this species. 

428 The Philippine Journal of Science i®^^ 

Between L. subnigra and D. oligolepia, there are differences 
in description which may be diagnostic of species. Thus, the 
paleae are described as "grosse dentatis" on the former, **integer- 
rimis" on the latter. On Brass 9062, they are mostly entire, 
but a few have marginal outgrowths. The indusium of the 
former is "glabrum," of the latter "in juventute minutissime 
glanduloso-puberulum." Some globose unicellular glands can be 
detected on the margin of the indusium of the Brass plant. I 
do not suppose that the differences are real, but have not been 
an authentic specimen of either species. 

25. L. CORIACEA (Brause) Copel., comb. nov. 

Dryopteris coriacea Brause, Engler's Jahrb. 56 (1920) 83. 

Known only by Ledermann's collections from the Sepik region, 
No. 10965, alt. 1,350 m (type), and No. 11291, alt. 1,300 m 
(var. elata). 

25a. L. KLOSSII Ridley 

L. Klossii Ridley, Trans. Linn. Soc. Bot. 9 (1916) 257. 

Carstenz Expedition, alt. 5,500 feet. 

This may be an older name for L. subnigra, but the description 
is inadequate for its recognition. 

L. erubescens (Wall.) Copel. is reported by Ridley in the 
paper just cited; it is otherwise unknown in New Guinea. 

20. h. STELLATO-PILOSA (Brause) Copel., comb. nov. 

Dryopteris stellato-pilosa Brause, Engler's Jahrb. 56 (1920) 96. 

Known only by the type collection, Schlechter 1685S, Mt. Gati, 
alt. 600 m. It is a small Leptogramma, having few elongate 
naked sori. 

n, L. WARIENSIS Copel. 

L. wariensis Copel., Genera (1946) 140. 

Dryopteris wariensis Copel., Philip. Jour. Sci. 6 C (1911) 73. 

Collected only once, — King s. n., Waria River, Papua. 

2S. L. TUBERCULATA (Ces.) Copel. 

L. tuherctdata (Ces.) Copel., Genera (1946) 140. 

Nephrodium tubercutatum Cesati, Atti Acad. Napoli 7 (1877) 29; 

see C. Chr., Dansk Bot. Arkiv 9 No. 3 (1937) 48. 
Dryopteris Schlechteri Brause, Engler*s Jahrb. 49 (1912) 16, /. 1 E, 
Dryopteris schizophylla v.A.v.R., Nova Guinea 14 (1924) 19. 

Brass 18656, alt. 700 m; 13885, alt. 150 m; Schlechter 16188, 
type of D. Schlechteri, Kelel, alt., 200 m; 16811, var. djamuensis 
Brause, Djamu, alt. 350 m, isotypes in Herb. Univ. Calif.; Lam 
1185, 1296, cotypes of D, schizophylla, of which the type, not 


Copeland: Aspidiaceae of New Guinea 429 

seen, is Lam 772. Remarkable for the absence of permanent 
paleae on the base of the stipe. 

29. L. NOTABILIS (Brause), Copel. 

L. notabilis (Brause) CopeL, Genera (1946) 139. 
Dryopteris notabilis Brause, Engler's Jahrb. 56 (1920) 91. 

Ledermann 11663 and H99l>, Sepik region, alt. 2,070 m, not 
seen. Brass 11283, alt. 2,200 m, is referable to this species. 
The stipe has a densely scaly base, conforming to Brause's 
German text, and is elsewhere aculeate, conforming to the Latin. 
The largest pinna is 30 cm long, 6.5 cm wide. 


30. L. DRYOPTEROIDEA (Brause) Copel. 

L. dryopteroidea (Brause) CopeL, Genera (1946) 138. 
Alsophila dryopteroidea Brause, Engler's Jahrb. 56 (1920) 70. 
Cyathea atrispora Domin, Acta Bohem. 9 (1930) 95. 
Dryopteris atrispora C. Chr., Brittonia 2 (1937) 296. 

Brass 10931, Lake Habbema, alt. 2,800 m, "Small tree-fern, 
trunk 70 cm tall, 6 cm in diameter; fronds few, ifc 1.5 m long." 
The type, not seen, is Ledermann 11897, Sepik region, alt. 2,070 
m. Brass' specimen conforms to the incomplete description. Its 
stipe is 65 cm long, sharply aculeate throughout, with black 
mostly curved spines, densely clothed in the lower part with 
narrow, chestnut-black, mostly appressed scales up to 1 cm long; 
sporangia naked. Like Brause, I have detected a sporangium 
with the annulus passing the pedicel; but it is exceptional 
practically all being of the polypodioid type. 


31. L. VARIEVESTITA (C. Chr.) Copel. 

L. varievestita (C. Chr.) CopeL, Genera (1946) 140. 

Dryopteris atrispora var. varievestita C. Chr., Brittonia 2 (1937) 296. 

Brass 4996, Mt Tafa, Central Division, Papua, alt, 2,400 m, 
in Herb. N. Y. Bot. Garden. As noted by Christensen, this is 
distinguished by the trichomes of the frond, and by not having 
veinlets connivent at the sinuses. The complete frond of L. 
dryopteroidea reveals more conspicuous differences, the stipe 
being much more spiny, and bearing on the lower part denser, 
narrower and much darker paleae. 

32. h, ALTA (Brause) Copel., comb. nov. 

Dryopteris alta Brause, Engler*s Jahrb. 56 (1920) 86. 
Ledermann 1U97, Sepik region, alt. 1,300 m; otherwise un- 

430 The Philippine Journal of Science i^^e 

33. L. QUADEIAURITA (Christ) Copel. 

L. quadriaurita (Christ) Copel., Genera (1946) 139. 
Dryopteris qtmdriaurita Christ, Philip. Jour. Sci. 2 C (1907) 209. 

King 220, Papua; Schlechter 17759. 


The published description should be amplified to show that 
the paleae are setulose, the lamina sparsely or deciduosly 
setulose above, the pinnae pinnate at the very base only, and 
the sporangia setulose. The King specimen is typical; that of 
Schlechter is nearly enough so (the pinnae are nowhere pinnate) . 

34. L. ENSIPINNA (Brause) Copel. 

L. ensipinna (Brause) Copel., Genera (1946) 138. 
Dryopteris ensipinna Brause, Engler^s Jahrb. 56 (1920) 84. 

Brass 12929, 1S362, alt. 1,200 and 850 m. I have not seen 
the type, Ledermann 12778, nor Ledermann 12533, var. acumi- 
nata, but Brass' collections fit the description. In minutiae, 
they are like L. qtmdriaurita, but have longer and narrower 
pinnae and more numerous segments. I suppose that the iden- 
tification of Schlechter 17759 is by Brause, and cannot but 
wonder that he failed to mention the resemblance. 



Dryopteris quadriquetra v.A.v.E., Nova Guinea 14 (1924) 16. 

Known only by the type collection, Lam 5i2, Mamberamo 
River, alt. 10 m. Not seen. Stipes glabrescent; fronds dimor- 
phic, subchartaceous. 

36. L. IMMERSA (Blnme) Moore 

L. immersa (Blume) Moore, Index (1857) 

Carr 1^299, Boridi, alt. 4,000 feet; previously reported. 
To Luzon and the Malay Peninsula ; reported from Polynesia 
to India. 

3T. L. KEYSSERIANA (Ros.) Copel. 

L. keysseriana (Ros.) Copel., Genera (1946) 139. 
Dryopteris keysseriana Ros., Fedde's Repert. 10 (1912) 333. 

Brass 12928, alt. 1,200 m. Described from Sattelberg, alt. 
1,600 m, Bamler 28 (1911) ; distributed as Rosenstock Fil. novog. 
exsicc. n. 121 1. Bamler. Remarkable for very long-caudate 
pinnae with oblique bases. Bamler 82c, received from Rosen- 
stock under a manuscript name, is a juvenile plant of this 


■^■^'^ Copeland: Aspidmceae of New Guinea 431 

38. L. OBLIQUATA (Mctt.) Copcl. 

L. obliquata (Mett.) Copel., Genera (1946) 139. 

Aspidium obliqtiatum Mett., Ann. Sc. Nat. IV 15 (1861) 75. 

Another species listed by report. Goadby 27, from New 
Britain, is similar but not identical. 
New Caledonia. 


Dryopteria Peekeli v.A.v.R., Bull, Dept. Agric. Ind. Neerl. No. 18 
(1908) 7. 

Known only by the type collection, Peekel 44, not seen. 
Evidently very similar to the preceding species. 

4d. L. GRACIL£SCENS (Blame) Hooker 

L. grcudlescenB (Blume) Hooker (as to the name), Jour. Bot. 9 
(1857) 338. 

Brass 12217, alt. 1,750 m. "On ground in a rain-forest ravine; 
rhizome purple when cut." New to New Guinea. 
To India and China. 

41. L. IAUTERBACHU (Branse) Copel. 

L, Lauterhaehii (Brause) Copel., Genera (1946) 139. 
Dryopteria Lauterhaehii Brause, Engler's Jahrb. 49 (1912) 18. 

Brass 12127, alt 1,800 m. The type is Schultze 273, Sepik 
region. Rhizome erect; stipes dark and densely scaly at base; 
scales sparse upward, caducous on rachis. 



Dryopteria stereophyla V.A.V.R., Nova Guinea 14 (1924) 17. 
Known only by the type, Lam 1785, Doorman-top, alt. 3,200 
m; not seen. 


DryopteHs rigidifolia v.A,v.R,, Nova Guinea 14 (1924) 18. 
Lam U81, 1560, 1562, 1890, Doorman Mountains, alt. 
1,420-1,450 m; not seen. This and the preceding species were 
to be expected in the Archbold Expedition collections, but I 
do not recognize them. 


Athyrium horizontale Ros., Nova Guinea 8 (1912) 722. 
Dryopteria horizontalia v.A,v.R., Bull. Buit. II No. XI (1913) 10. 

Hellwig-Gebirge, alt. 1,350-1,600 m, v. Roemer 1136; not 
seen. Except that it is described as "subcoriaceo-chartaceous," 
this would seem very near to L. gracilescens. 

10832 5 

432 The Philippine Journal of Science ^^*^ 

45. L. CALCARATA (Blume) Moore 

L. calcarata (Blume) Moore, Index (1858) 87. 

Known from New Guinea by Ledermann 117^0, det. Brause. 
Java to India, common and plastic; not known in the Philip- 

46. L. FULGENS (Brause) Copel., comb. nov. 

Dryopteris fulgens Brause, Engler's Jahrb. 56 (1920) 89. 

Known only by the type collection, Ledermann llOOi, Hun- 
stein Peak, alt. 1,300 m ; not seen. A large relative of L. falea- 

47. L. SUBDIMORPHA Copel. Plate 20. 

L. subdimorpha CopeL, Genera (1946) 140. 

Dryopteris subdimorpha CopeL, Univ. Calif. Publ. Bot. 18 (1942) 220. 

L. rhizomate horizontal 1 cm crasso, basibusque stipitum 
confertorum paleis fuscis integris plerisque 2-3 nrni longis lan- 
ceolatis nonnulis 5 mm longis linearibus vestitis ; stipite f rondis 
sterilis ca. 25 cm alto, f rondis fertliis altiore, brunneo, paleis 
parvis linearibus sparso, sursum rhachique pubescentibus parce 
paleolatis; lamina sterile 30 cm vel ultra longa, 20-25 cm lata, 
subcoriacea, faciebus glabris, apice acuminata pinnatifida, basi 
truncata; pinnis infimis subdeflexis 13 cm longis 3.5 cm latis, 
pedicellis 1 cm longis protensis, apicibus integris breviter acu- 
minatis, basibus rotundatis, segmento infimo acroscopico obso- 
leto, basiscopico abbreviato, alibi ad alam 5 mm latam costae 
suboblique pinnatifidis, segmentis 6 mm latis obtusis integris 
margine leviter cartilagineo-inerassatis ; costis utraque facie 
setul(^is etiam infeme paleis paucis minutis obscuris sparsis; 
costulis nudis, venis ca. 10-paribus, infimis ad vel infra sinus 
conniventibus, infima basiscopica a costula remota e costa ^re- 
diente; lamina fertile minore laxiore; soris medialibus, exindu- 
siatis sporangiis nudis. 

Dutch New Guinea: 2 km SW. of Bemhard Camp, Idenburg 
River, alt 700 m, Brass 1S666; "rain-forest undergrowth; few 
tufts on crests of ridges." 

Near enough to be conspecific is Brass 5155, from Mafulu, 
Papua, alt. 1,250 m, in Herb. N. Y. Bot. Garden, identified by 
Christensen as D. crassifolia var. purpureo-lilacina C. Chr. Its 
pinnae are more numerous, smaller and in particular narrower, 
at most 23 mm wide, and the lowest are somewhat deflexed. 
The type of L, subdimorpha is mature, and I can only say of 
it that no indusia are present; but Brass 5155 has an immature 
fertile frond, and sori without any mature sporangia are exin- 

^"^'^ Copeland: Aspidiuceae of New Guinea 433 

dusiate, I do not adopt Christensen's varietal name, because 
uncertain of identiy with the Bornean varietal type; also, it 
is too long a name. 

The most immediate affinity seems to be to L. motleyana 
(Hooker) Copel., whether that be a distinct species, or, as 
Ching, Bull. Fan. Bot. 6 (1936) 286, regards it, a variety of 
L. crassifolm. The pinnae of L. subdimorpha are more deeply 
cut. There is some uncertainty as to the indusia of L. crassi- 
folia and L. motleyana. My specimens of L. crassifolia suggest 
no dimorphism. 


L. falcatipinnula Copel., Genera (1946) 138. 

Dryopteris falcatipinnula Copel., Philip. Jour. Sci. 6 C (1911) 74. 

King Hi, Papua, in lowlands (1908). We have a larger and 
more perfect specimen than the type, — Rosenstock Fil. novog, 
exsicc. n. 199, supposed to be a part of the original collection, 
but dated 1911. 

49. L ECHINATA (Mett.) Copel. 

L. echinata (Mett.) Copel., Genera (1946) 138. 

Aspidium echinatum Mett., Ann. Lugd. Bat. 1 (1864) 230. 

Dryopteris echinata C. Chr. See Gardens' Bull. 4 (1929) 386. 

Listed by Brause, Engler's Jahrb. 56 (1920) 81. I have a New 
Guinea specimen collected by Zahn, Sattelberg, alt. 900 m, 
received with this name, but misidentified. 

50. L. BELENSIS Copel. Plate 21. 

L. belensis Copel., Genera (1946) 138. 

Dryopteris belensis Copel., Univ. Calif. Publ. Bot. 18 (1942) 220. 

L. gregis L. crassifoliae, rhizomate erecto crasso, inter bases 
stipitum radicumque paleis fuscis ovatis 1.5 mm longis dense 
setulif eris vestito ; stipitibus caespitosis, 40 cm altis, fuscis, basi 
paleis parvis sat dense vestitis et hie solummodo tuberculatis, 
sursum rhachibusque paleis lanceolatis et linearibus ciliatis 1-2.5 
mm longis sparsis et minute setuliferis, rhachibus infra bases 
pinnarum glandulif eris ; lamina 45 cm longa, 25 cm lata, acu- 
minata, bipinnatifida, fusca, tenuiter coriaceae; pinnis typi 
13-paribus, sessilibus, infimis vix abbreviatis haud deflexis, 
sequentibus 15 cm longis, 3 cm latis, caudato-acuminatis, basi 
rotundatis, ad alam 2 mm latam costae pinnatifidis, costis 
superne sulcatis utraque facie setulis brunneis haud densis 
obsitis; segmentis erecto-patentibus plus minus falcatis, infimis 
plerumque modo reductis, sequentibus 2 cm longis, basi 5 mm 
latis, acutis, integris, decidue ciliolatis, superne glabris infeme 

434 The Philippine Journal of Science ^^^^ 

ad costulas sparse et decidue setuliferis; venis usque ad 18- 
paribus, simplicibus, infimis ad vel infra sinus conniventibus 
vix anastomosantibus, venula infima basiscopica a costula remota 
e costa oriente; soris costularibus, indusiis parvis setulosis, 

Dutch New Guinea: Bele River, alt. 2,200 m. Brass 11509, 
"on a limestone cliff in forest; not common;" also, No. 11327, 
same place, an older, somewhat larger specimen, the indusia 
gone, the sori plainly marked by minute depressions on the 
upper surface. 

This species might be responsible for the report of L. echinata 
in New Guinea, being like that species in connivent veins and 
slightly scaly rachis. It is more like L. Hallieri, but has less 
setulose costae and costules. Its acute segments are peculiar 
in the group. 

51. L. COSTUUBSORA Copel. 

L, costulisora Copel., Genera (1946) 138. 

DryopteHs basisora Copel., Philip. Jour. Sci. 6 C (1911) 73, non 
Christ (1909). 

Brass 12^65, alt. 2,150 m. The type is King BOh, from Papua, 
distributed as Rosenstock Fil. novog, exsicc. n, 198, but mis- 
dated 1911. Keysser 23p, Sattelberg Hinterland, received indet. 
from Dr. Rosenstock, is also this species. This is a member 
of the group of L. crassifolia, its lowest veins connivent at the 
sinus, with a raised (on the lower surface) line running from 
the sinus between them. Rhizome ;stout, short-creeping, beset 
with firm, brown densely setuliferous paleae 5 mm long; indusia 
dark, naked, firm and persistent. L. wariensis, to which Chris- 
tensen, Suppl. I 30. thought this might be reduced, is not a 
near relative; it is exindusiate, and the lowest veinlets run to 
the margin above the sinus. 


52. I.. CHUIMYDOPHORA (Bos.) Copel. 

L. cMamydophara (Eos.) Copel., Genera (1946) 138. 

Dryopteris chlamydophora Eos. C. Chr,, Gardens' Bull. 4 (1929) 384. 

Brass 3808, Papua, alt. 500 m; see Brittonia 2 (1937) 297. 
Celebes to Burma. 

53. Ii. OPHIUEil Copel, Plate 22. 

L. ophiura Copel., Genera (1946) 139. 

Dryopteris ophiura Copel., Univ. Calif. Publ. Bot. 18 (1942) 220. 

L. rhizomate erecto, valido, paleis fuscis minute setuliferis 
4 mm longis dense vestito; stipitibus approximatis, 15 cm altis, 

'^'^'^ Copeland: Aspidiaceae of New Guinea 435 

1 mm crassis, basi imo paleatis, alibi rhachibusque minute pallida 
pubescentibus stramineis; lamina 20 cm longa, ca. 10 cm 
lata, acuminata, basi truncata, coriacea, costis superne minute 
antrorso-strigosis exceptis glabra, bipinnatifida ; pinnis ca. 
7-paribus, infimis vix reductis haud deflexis, superioribus 1-2- 
paribus sat abrupte diminutis et integrescentibus, medialibus 
ca. 6 cm longis, 14 nun latis, in caudas 2 cm longas rectas 
f alcatas vel sinuosis protractis, subsessilibus ; segmentis infimis 
reductis rarius liberis, infimo basiscopico interdum tantum ves- 
tigiale; segmentis medialibus ca. 8 mm longis, 2-3 mm latis, 
subfalcatis, integris margine subdeflexis, obtusis vel oblique 
acutis; venulis usque ad 10-paribus, immersis, infimis supra 
sinus terminantibus ; soris inframedialibus, indusiis coriaceis 
persistentibus glabris. 

Dutch New Guinea : 18 km SW. of Bernhard Camp, Idenburg 
River, alt. 2,150 m. Brass 12^71, small clumps on open rock-slide. 
There is an evident, but not salient, gland on the rachis under 
the insertion of each pinna. 

54. L. CRASSA Copel. Plate 23. 

L. crassa Copel., Genera (1946) 138. 

Dryopteris crassa Copel., Univ. Calif. Publ. Bot. 18 (1942) 220. 

L. rhizomate repente 1 cm crasso, paleis fuscis 1-2 mm longis 
lanceolatis minutissime setulosis vestito ; stipitibus approximatis, 
60 cm altis, 3 mm crassis, brunneis, basi imo paleatis alibi 
glabris; lamina 20-30 cm longa, 15-20 cm lata, acuminata, 
basi truncata, rhachi et deorsum costis minute purpureo-stri- 
gosis, alibi glabra, rigide coriacea, bipinnatifida; pinnis infimis 
vix diminutis haud deflexis, segmentis infimis reductis; pinnis 
medialibus frondis maximae 11 cm longis, basi truncatis 2 cm 
latis, caudato-acuminatis ; segmentis 1 cm longis, 3-3.5 mm 
latis, plerisque subfalcatis, obtusis, margine deflexis, integris; 
venulis ca. 12-paribus, conspicuis, infimis pauUo supra sinus 
terminantibus; soris subcostularibus, indusiis obscuris nudis 
fugacibus. Specie praecedente (L. ophiura) differt: rhachi cos- 
tisque infeme glabris, indusia deciduis. 

Dutch New Guinea: 9 km N. E. of Lake Habbema, alt. 
2,650 m, Brass 10878, type; ibidem No. 109Si. 

Because it grew on a rock-slide, I do not trust the erect 
rhizome of L. ophiura as a fixed character. It and L. crassa 
rook much alike. The darker stipe and rachis, less caudate tips 
of the pinnae, and slightly greater stature of L. crassa may not 
be essential differences, but the naked axes and transient indusia 

436 The Philippine Journal of Science i^*» 

should be diagnostic. I am loath to describe these two coriaceous 
specie while two others, D. stereophylla and D. rigidifolia, from 
the same region, are unknown to me, but am unable to reconcile 
the difference in description. 

S5. L. WANTOTENSIS Copel. Plate t4. 

L. wantotensis Copel., Genera (1946) 140. 

Dryopteris wantotensis Copel., Univ. Calif. Publ. Bot. 18 (1942) 220. 

Clemens 110 IS bis, Morobe; known by the one collection, 

SC. h, NEPHROLEPIOIDI^ (C. Chr.) Copel. 

L, nephroteptoides (C. Chr.) Copel., Genera (1946) 189. 
Dryopteris nephroteptoides C. Chr., Brittonia 2 (1937) 295, Fig. 1 
(p. 268) c, d. 

Known only by the type collection, Brass 585^, Maf ulu, Papua, 
alt. 1,700 m. 

57. L. HUNSTEINIANA (Brause) Copel. 

L. hunsteiniana (Braxise) Copel., Genera (1946) 139. 
Dryopteris hunsteiniana Brause, Engler's Jahrb. 56 (1920) 79. 

Brass 12872, alt. 1,500 m, "on shaded bank of stream; fronds 
glaucous beneath." The type is from Hunstein Peak, alt. 1,350 
m, not seen but unmistakable from the description. Brass' 
plant has the lowest pinnae enlarged and incised, the second 
pair like the segments above, only free, and remote by about 
5 mm. 


58. L. SETOSA Presl 

L. setosa Presl, Epim. (1849) 40. 

Cheilanthes setigera Blume, Ennm. (1828) 138. 

Lastrea setigera Bedd., Ferns Brit. India, Corr. II (1870), nan 

Dryopteris setigera O. K., Kev. Gen. PL II (1891) 813. 

Common at minor altitudes. Polynesia to India. 

59. L. ARMATA Copel. Plate 25. 

L. armata Copel., Genera (1946) 138. 

Dryopteris sjnnosa Copel., Univ. Calif. Publ. Bot. 18 (1942) 219. 
non Lastrea spinosa Newman. 

L. leucolepidi similis, paleis basalibus castanesis ovatis glabris 
axibus minoribus minute puberulis distincta; stipite 75 cm alto 
basi 1 cm sursum (sicco) 4 mm crasso, stramineo, deorsum 
paleis castaneis 1-2 cm longis 4-5 mm latis acutis vel acuminatis 
glabris baud ciliatis vestito, sursum rhachique sparse minutis- 
sime puberulis ubique spinis 0.5-1.0 mm longis horridis; lamina 

'^'^' ^ Copeland: Aspidiaceae of New Guinea 437 

80 cm alta, ovata, acuminata, tripinnata, firme papyracea, f us- 
cescente, supeme (etiam in sulcis axium) glabra, rhachibus 
infeme spinulis sursum minutis armatis, minute obscure debi- 
liter pubescentibus ; pinnis infimis 15 cm, medialibus 25 cm 
longis, 6 cm latis, acuminatis, breviter pedicellatis; pinnulis 
subsessilibus, recte patentibus, acuminatis, basi 1 cm latis pin- 
natis, alibi fere ad costam pinnatifidis, segmentis et pinnulis 
secundariis usque ad 12-paribus infra apicem inciso-serratum, 
majoribus crenato-dentatis; soris laminam omnino tegentibus, 
indusiis nullis. 

Dutch New Guinea: 9 km N. E. of Lake Habbema, alt. 2,800 
m, Brass 10751. "Gregarious in young second growth on a 
native clearing." This is not the place to expect a new species ; 
it is a typical place for L. setosa and L. leucolepis, both of which 
enter abandoned clearings in the Philippines like weeds. This 
species is like them in dissection, and thus in general aspect, 
but most distinct in its nakedness. 

60. L. LEUCOLEPIS Presl 

L. leucolepis Presl, Epim. (1849) 39. 

Brass 12^62, alt. 2,150 m; Carr 12SS9, Ronna, alt. 1,400 feet; 
probably common. In naming Carr's specimen, Alston has 
agreed with others who call this Dryopteris uUginosa (Kunze) 
C. Chr. ; for some synonyms under that name, see Christensen, 
Suppl. Ill 100. Maxon, Proc. Biol. Soc. Washington 36 (1923) 
170, has interpreted Aspidium uUginosum as D. setigera, L, 
setosa of this enumeration. Backer and Posthumus, Varenflora 
voor Java, p. 42, treat D. leucolepis as a synonym of D. uliginosa, 
Christensen, Suppl III 90, Dansk Bot. Arkiv 9 No. 3 (1937) 44, 
treats D. leucolepis as distinct, and cites New Guinea synonyms, 
Cheilanthes gigantea Ces., Polypodium cheilanthoides Baker, 
and Dryopteris brunneo-villosa C. Chr. The original description 
of A. uUginosum Kunze does not show whether it is L. setosa 
or L. leucolepis; it is almost certainly one or the other, and 
its name has priority. Unable to settle the question, I follow 
Maxon; but compare still Christensen, Engler's Jahrb. 66 
(1933) 46. 

Polynesia to India and Japan. 

24. CURRANIA Copeland 
C. OYAMENSIS (Baker) Copel. 

C. oyamensis (Baker) CopeL, Philip. Jour. Sci. 6 C (1911) 147. 
Dryopteris genuflexa Ros., Fedde's Repert. 12 (1913) 175. 

438 The Philippine Journal of Science *^^^ 

Keysser B 56 Bolan, alt. 2,400-3,000 m ; isotype of D, genuflexa 
in Herb. Univ. Calif. 
Japan; China. 


Hort. Berol. (1833) 128; Ching, Bull. Fan Bot. 8 (1938) 162; Cope- 
land, Genera (1946) 140. 

Cyclosorus is so intimately related to Lastrea that its separa- 
tion is justified only by convenience, both being very large 
genera. New Guinea may be regarded as the center of evolution 
of Cyclosorus, 

Key to the species 
Sori round. 
Only one pair of veinlets anastomosing. 
Fronds under 5 cm broad. 

Frond practically naked 1. C roemerianus 

Frond strigose. 

Fronds uniform 2. C Warburgii 

Fertile fronds contracted 66. C. canescens var. 

Frond finely pubescent 3. C. obtusifolius 

Fronds 7-10 cm broad. 

Pinnae subentire 8. C. mutabUis 

Pinnae sharply serrate 9. C. cataractorum 

Pinnae lobed % to costa 4. C. adenostegms 

Pinnae incised over half-way to costa. 5. Dryopteris repandula 

Fronds comparatively large. 
Margin of pinnae entire to serrate. 
Indusium naked. 

Fronds uniform 9. C Cataractorum 

Fronds dimorphic 6. C. dimorphus 

Indusium setulose 7. Dryopteris sepikensis 

Pinnae pectinate. 
Basal pinnae not reduced. 
Veins setose beneath. 
Pinnae under 15 cm long. 

Caudex erect 11. C albociliatus 

Rhizome creeping 12. C parasiticus 

Pinnae 25 cm long 13. C distinctus 

Veins naked beneath 14. C. alatellus 

Basal pinnae gradually reduced. 
Segments 2-2.5 mm wide. 

Frond about 50 cm tall 15. Dryopteris perpili- 


Frond over 1 meter tall 16. Dryopteris hispiduli- 

Segments 4-5 mm wide. 
Glaucous beneath. 
Lamina membranaceous 17. C. austerus 


Copeland: Aspidmceae of New Guinea 


Lamina chartaceous 18. C. farinosus 

Not glaucous. 

Lamina setulose 19. Dryopteris hirto-pi- 


Lamina naked 42. C. truncatus 

Basal pinnae abruptly reduced. 
Lamina setulose above. 

Yellowish-gray beneath 18. C. farinosus 

Greenish beneath. 
Saudex erect. 

Veins naked above 20. C. suprastrigosus 

Veins setose above 21. C. vestigiatus 

Rhizome creeping 22. C. obUmgus 

Lamina naked above. 

Lamina over 1 meter long 23. C. superbus 

Lamina smaller 24. C. heterocarpus 

More than the lowest veinlets anastomosing. 
Basal pinnae abruptly reduced. 
Margin cut at least one-fourth costa. 
Lamina naked above. 

Sori strictly costular 25. C. supraspinigerus 

Sori inframedial or medial. 
Nether surface setulose. 

Pinnae alternate 26. C. paraphysatua 

Pinnae opposite 27. C. paripinnatus 

Nether surface naked. 
Pinnae opposite. 

Pinnae dilated at base 28. C. deltipterus 

Pinnae not dilated 29. C. subappendiculatus 

Pinnae alternate 30. C. sogerensis 

Sori extramedial 31. C. extenstis 

Lamnia setulose above. 

Veins densely setose beneath 32. C. megaphylUndes 

Veins almost naked beneath 32. A. C, sp. 

Margin entire or shallowly toothed. 
Pinnae numerous. 
Subcoriaceous or thinner. 

Pinnae opposite 29. C. subappendiculatus 

Pinnae alternate. 

Setose above 33. C. Atasripii 

Glabrous or nearly so. 

Veins 3 or 4 pairs 34. Dryopteris tamiensis 

Veins about 6 pairs. 

Pinnae truncate at base 35. c. arfakianus 

Pinnae cuneate 30. C. gregarius 

Coriaceous 37. C. unitus 

Pinnae few — 3-6 pairs 61. C, micans 

Basal pinnae gradually reduced. 
Upper surface naked or glabrescent. 
Pinnae 15 cm long or longer. 

440 The Philippine Journal of Science ^^*® 

Pinnae lobed. 

Indusia hairy 40. Dryopteris piloso- 


Indusia not hairy. 

Pinnae cut one-fourth to costa 41. C megaphyllus 

More deeply cut 42. C. truTicatus 

Pinnae crenate to entire 35. C arfakianus 

Pinnae hardly exceeding 10 cm. 
Apparently exindusiate. 

Paleae naked 43. C. rigidus 

Paleae hairy 44. 0. multiauriculatus 

Evidently indusiate. 
Pinnae under 15 mm wide. 

Pinnae cuneate 53. C riparius 

Truncate at base. 
Sori about medial. 

Rhizome creeping 38. C aridiis 

Rhizome ^ erect 45. C protecttis 

Sori subcostular 39. C Archboldii 

(Pinnae 2 cm wide) 
Pinnae 2 cm wide. 

Pinnae gradually narrowed 46. C ainboinensis 

Pinnae abruptly narrowed 47. C terrestris 

Upper surface setulose. 

Pinnae opposite 29. C suhappendiculattis 

Pinnae alternate. 
Upper surface sparsely hairy. 

Pinnae up to 1 cm wide 39. C Archboldii 

Pinnae 15 mm wide 48. C confertus 

Upper surface freely hairy. 

Pinnae 15 mm wide 49. C. dichrotrichus 

Pinnae up to 11 mm wide. 
Auricles on stipe remote. 

Pinnae linear-lanceolate 50. C hispidulus 

Pinnae broadly lanceolate 51. C, albosetosus 

Auricles crowded 52. C strigosissimus 

Basal pinnae not reduced. 

Pinnae over 30 cm long 54. C. imponens 

Pinnae much shorter. 
Pinnae 1^3S than 2 cm wide. 

Sori restricted to lobes 55. C, interruptiis 

Sori not so restricted. 

Lamina setose beneath 56. C invisus 

Lamina naked. 

Costae naked 57. C. gongylodes 

Costae setulose 10. C. aquatilis 

Sterile pinnae over 2 cm wide. 

Lamina glandulose beneath 58. C, glanduloaus 

Lamina not glandulose. 

Pinnae abruptly caudate 59. C. cuspidatus 

Pinnae not caudate 60. C. urophyllus 

'^^'^ Copeland: Aspidiaceae of New Guinea 441 

Sori elongate, or basal ones confluent. 
Pinnae numerous. 
Base of frond gradually narrowed. 

Base of pinna auriculate 62. C. hastato-pinnatus 

Base of pinna cuneate-rounded 63. C morobensis 

Base of frond truncate. (Ampelopteris) 

Pinnae about 5 pairs, all small 67. C lanceoltia 

Pinnae 1-3 pairs, fairly large. 

Margin entire or nearly so 64. C. triphylltis 

Margin lobed 65. C. pentaphyllus 

Frond simple or with few reduced pinnae. 

Body of frond inciso-lobate 66. C. eaneseena 

Body of frond more or less entire 68, C. heccwnanus 

1. C. ROEMERIANUS (Ros.) Copel. 

C. roemerianus (Ros.) Copel., Genera (1946) 139. 
Dryopteris roemeriana Ros., Nova Guinea 8 (1912) 723. 

Brass 1U97 (?), Bele River, alt. 2,200 m, "abundant on 
limestone cliffs in forest." The type, not seen, is von Roemer 
1025, in mountain forest, alt. 1,350 m, also in Dutch New 
Guinea. Brass' plant presents inconsistencies with the descrip- 
tion, most essentially in having one perfectly regular row of 
sori on each side of the costa, and in having elongate fronds, 
about 30 cm long (length not stated by Rosenstock, but shortness 
indicated by stated shape) ; but there are so many points of 
agreement that I am unwilling to describe it as distinct from the 
unseen species of the same region. The veinlets do not anas- 
tomose on the basiscopic side of the pinnae, not anywhere on 
the smaller pinnae; but on the acroscopic side of the larger 
ones, the lowest veinlets of seceral veins regularly anastomose, 
with "radio calloso in sinum excurrente." The probable affinity 
is not to Cyclosoms, but to Lastrea nepKrolepioides and L. 
petrophila. Endemic. 

2. C. WARBURGII (Kuhn and Christ) Wagner and Grether 

C Warburgii (Kxihn and Christ) Wagner and Grether. 
Aspidium Warburgii Kuhn and Christ, Monsunia 1 (1900) 81. 

Bamler 131 (1914) Sattelberg (type locality of C Warburgii) , 
alt. 400 m, received as Dryopteris dehilis (Mett.) C. Chr., is 
more exactly C. Warburgii, distinguished by persistent, setose 
indusia. Clemens 6522, Morobe, alt. 1,700 feet, has broader, 
l^s densely hirsute pinnae. 

Admiralty Ids.; Guam (as Dryopteris depauperata Copel.). 

442 The Philippine Journal of Science 


3. C. OBTUSIFOUUS (Ros.) Copel. 

C. obtusifolius (Ros.) Copel., Genera (1946) 143. 
Dryopteris obtusifoUa Ros., Fedde's Repert. 10 (1912) 336. 
Known only by the type collection, Bamter 115, Sattelberg, 
alt 700 m; isotype in Herb. Univ. Calif. 

4. C. ADENOSTEGIUS Copel. Plat« «»»• 

C. adenostegius Copel., Genera (1946) 142. 

Dryopteris adenostegia Copel., Univ. Calif. Publ. Bot. 18 (1942) 220. 

C. rhizomate brevi-repente 5 mm crasso; stipitibus approx- 
imatis, frondis sterilis 10 cm, fertilis 30 cm altis, fuscis, basi 
paleis paucis fuscis minutis praeditis,* alibi cinereo-velutinis vel 
sursum rhachibusque potius dense setulosis; lamina ca. 30 cm 
longa, 8-9 cm lata, acmninata, basi angusta; pinnis inferioribus 
deflexis paucis sensim diminutis, infimis 8 mm longis 5 mm 
latis oblique deltoideis; pinnis medialibus 4-4.5 cm longis, 12 
mm latis, obtusis vel acutis, basi truncatis inconspicue auricu- 
latis, ca. one-third ad costam incisis, segmentis 2-2.5 mm latis, 
subcoriaceis, costis sparse costulisque sparsius strigosis, venis 
superne sparsissime setosis, facie superiore laminae sed cons- 
picue setulosa; venis ca. 5-paribus, infimis anastomosantibus, 
sequentibus 1-paribus ad sinus conniventibus ; soris inf ramedia- 
libus, indusiis glandulis globosis aurantiacis cinctis, sporangiis 

Dutch New Guinea: 9 km N.-E. of Lake Habbema, alt. 2,800 
m, terrestrial in forest. Brass 10282; type in Gray Herbarium. 

This may resemble Dryopteris repandula V.A.V.R., the descrip- 
tion of which is more elaborate than clear; it (C adenostegius) 
seems to be distinguished by absence of paleae above the base 
of the stipe, less deeply incised pinnae, inframedial sori, and 
gland-bordered indusia. 


Dryopteris repandula V.A.V.R., Nova Guinea 14 (1924) 20. 

Known only by the type, JLam 1058, Mamberamo River, alt. 
90 m; not seen. 

6. C. DIMORPHUS (Brause) Copel.» comb. nov. 

Dryopteris dimorpha Brause, Engler's Jahrb. 56 (1920) 100. 

Known positively only by the type, Ledermann 12622, Sepik 
region, alt. 1,400-1,500 m, not seen. Carr 15531, sent to me as 
this species, has meniscioid venation, which cannot be reconciled 
with Brause's "Nervis . . . infimis anastomosantibus;" and 
there are other discrepancies. 


Copeland: Aspidiaceae of New Guinea 443 

From C. alatellus, which likewise is commonly dimorphic, 

C. dimorpha seems to be distinguished by very reduced lower 
pinnae, and by normal pinnae less contracted toward the rachis. 


Dryopteris sepikensis Brause, Engler's Jahrb. 56 (1920) 101. 

Known by the original collections, Ledermann 12053 and 
11183, Sepik region, alt. 2,070 and 1,300 m, not seen. Judging 
by description only, this seems suspiciously like the preceding 
species. Christensen, Suppl. Ill 97, reduces it to D. arfaJdoma 
(No. 35), which is hard to reconcile with Brause's description. 

8. C. MUTABIUS (Brause) Copel. 

C. mutabilis (Brause) Copel., Genera (1946) 143. 
Dryopteris mutabilis Brause, Engler's Jahrb. 56 (1920) 97. 

Brass 1S0A7, alt. 850 m, on flood-swept river bank. The 
type is Ledermann 97^5, Kaiserin-Augusta River, alt. 200-400 
m, isotype in Herb. Univ. Calif. The narrowness of the pinnae 
is responsible for the mostly free veinlets. 

9. C. CATARACTORUM Wagner and Grefher 

C. cataractorum Wagner and Grether, Pterid. 

Rook Island, Bamler R 14, reported by Rosenstock as ''prope 

D. aquatilem Copel.", which is correct as far as it goes. It is 
distinguished from C, aqttatilis by sharply toothed pinnae more 
narrowly cuneate at the base, and by a single pair of anastomos- 
ing veinlets. 

Admiralty Ids., along lowland streams. 

10. C. AQUATILIS Copel. 

C. aqtuitilis Copel., Genera (1946) 142. 
Dryopteris aquatilis Copel., Philip. Jour. Sci. 6 C (1911) 75. 

Known only by the type collection. King 182, Papua, below 
the flood level of creeks. 

C. Warburgii, C. mutabilis, C. aqvxitilis and C. riparia (No. 
53) are all stream-bank plants, similar in adaptation to this 
habitat, and all probably related. 

11. C. ALBOCIUATUS Copel. 

C. alhodliatus Copel., Genera (1946) 142. 

Dryopteris albodliata Copd., Jour. Arnold Arb, 10 (1920) 177. 

The type is Brass 566, Basiatabu, Papua, alt. 1,500 feet. To 
this species I now refer King 405, from Loane; it is somewhat 
larger than the type, with broader instead of more numerous 
segments, and with the same mixture of pale bristles. 


444 The Philippine Journal of Science ^^*^ 

12. C. PARASITICUS (L.) FarweU 

C. parcmticus (L.) Farwell, Amer. Midland Nat. 12 (1931) 258. 

Carr 11645, Veiya, Papua, at sea level, received as Dryopteris 
didymosora (Parish) C. Chr. This identification seems correct, 
but I can see no detail of difference from the plant I have from 
Fokien and Tonkin, which is C. parasiticus as of late strictly 
construed. The rhizome is long-creeping. This species, or C. 
amboinensis, may be responsible for the listing of Dryopteris 
mollis (Jacq.) Hieron. as a New Guinea species. 

Westward to India. 

13. C. DISTINCTUS Copel. Plate 26. 

C distincius Copel., Genera (1946) 142. 

Dryopteris distincta Copel., Univ. Calif. Publ. Bot. 18 (1942) 220. 

C. rhizomate ignoto; stipite deorsum fusco, paleis fuscis 
membranaceis glabris usque ad 12 mm longis basi 1-1.5 mm 
latis vestito, sursum rhachique brunneis sparsius et decidue 
paleatis, paleis dejectis spinulis parvis nigris sparsis; lamina 
grande, herbacea, bipinnatifida, apice breviter acuminata, basi 
late truncata; pinnis infimis horizontalibus ca. 25 cm longis, 
3 cm latis, acuminatis, basi modo angustatis subsessilibus, 
sequentibus sessilibus basi utroque latere dilatatis, aliter cum 
basalibus conformibus, ad alam 1.5-2.0 mm latam costae pecti- 
natis, segmentis permultis patentibus subfalcatis obtusis vel 
subacutis integris usque ad 2 cm longis 2.5-3.0 mm latis ; costa 
supeme strigosa, inferne costulisque venisque setulosis, facie 
inferiore laminae minutissime setulosa; venis ca. 17-paribus, 
infimis anastomosantibus, sequentibus non conniventibus, soris 
inframedialibus, indusiis nullis vel vestigialibus, sporangiis 

New Guinea, present mandated territory : Wareo, alt. 150-180 
m, Bamler W 11 (March, 1914), in Herb. Univ. Calif. No. 
3 9 mi, type; alt. 600 m, Bamler, Rosenstock Fil, novog. exsicc. 
n. 248. 

This is Dryopteris longissima (Brack.) var. novoguineensis 
Ros., Hedwigia 56 (1915) 351, the type collection of which was 
published as Bamler 182; it was presumably the collection dis- 
tributed as Fil. novog. exsicc. n. 248. As a complete description 
was needed, and the varietal name could not serve as specific, 
I chose as the type our more perfect specimen. I have observed 
before. Bull. B. P. Bishop Museum No. 93 (1932) 38, that 
C longissimus is endemic in Tahiti, and this New Guinea fern 
is far too distinct to justify their association. As they are not 
particularly related, the effect of treating the New Guinea species 

''^' * Copeland: Aspidiaceae of New Guinea 445 

as a variety is to present false evidence of floristic affinity 
between New Guinea and Polynesia. 

14. C. ALATELLUS (Christ) Copel. 

C. alatellus (Christ) Copel., Genera (1946) 142. 
Nephrodium and Aspidium alatellum Christ (1901). 
Dryopteris stenobasis C, Chr., Index (1905) 294; older synonyms here. 

D. logavemis Ros., Fedde's Eepett. 10 (1912) 332. 

? D, pseicdostenobasis Copel., Jour. Arnold Arb. 10 (1929) 176. 

First collected in New Guinea by Weinland, but described from 
the Schuman and Lauterbach collection. In hand are : Schlechter 
18038^ Finisterre Mountains, alt. 800 m; Bamler 84 and S5, the 
latter being Rosenstock Fil. novog. exsicc. n. 78,'D, logavensis, 
the description of which is based on No. 35; Zahn ss. nn., 
Sattelberg, alt. 900 m, purporting to be determined by Christ 
as Nephrodium echinatum and N. pteroides, but without re- 
semblance to either; Bamler, Rosenstock Fil. novog. exsicc. n, 
230, also from Logaueng, and recognized as D. alatella; Brass 
5U0, Mafulu, alt. 1,250 m. 

The fertile frond and pinnae of this species may or may not 
be contracted. Christ noted the dimorphism of the Weinland 
collection. Rosenstock distinguished his D. logavensis by its 
not being dimorphic, — ^which is most remarkable, because Bamler 
3U) Rosenstock's forma minor, is the contracted form. Both 
are present on the Zahn collection received as D. pteroides. 
Both are also present on my isotype of D. stenobasis, Cuming 
327; less distinct on Ramos, Bu. ScL 17600, from Samar; and 
as somewhat different pinnae on the same frond of Elmer 13555 
A, from Mindanao. It is a variable species in other respects 
also, — in the width of its unconstricted segments, and in the 
indusium, which is more or less fugacious. Because I now 
recognize this as a variable species, I mistrust the distinctness 
of D. pseudostenobasis, which has no trace of an indusium, and 
has one or two (perhaps casually) reduced lower pinnae. 

Central and Southern Philippines; Celebes. 


Dryopteris perpilifera v.A.v.R., Bull. Jard. Bot. Buitenzorg No. XI 
(1913) 12. 

Known only by the original collection, Lauterbach 68, from 


Dryopteris hispiduUformis C. Chr., Suppl. Ill (1934) 88. 

D. hispida Brause, Engler's Jahrb. 56 (1920) 102, non O. K. 

446 The Philippine Jounml of Science ^^^^ 

Authentically known only by the original collections, Leder- 
mann 11758 and 11709, Sepik region, alt. 2,070 m. See No. 81 
A of this list. 

17. C. AVStFERUS O^rause) Copel., comb. nov. 

Dryopteris austera Brause, Engler's Jahrb. 56 (1920) 108. 

Known only by the type collection, Ledermann 11750, Sepik 
region, alt. 2,070 m. Basal pinnae gradually reduced to rudi- 
ments; sori exindusiate. 

18. C. FARIKOSUS (Brause) Copel., comb. nov. 

Dryopteris farinosa Brause, Engler's Jahrb. 56 (1920) 111. 

Known only by the type collection, Ledermann 1210S, Sepik 
region, alt. 2,070 m. Not too evidently distinct from C austerus 
from tiie same locality, which is described as larger and thinner. 


Dryopteris hirto-pilosa Ros., Meded. Rijks Herb. No. 31 (1917) 7. 

Carr 111 IS, 11727, Papua, near sea level, det. Alston. 

The type is Merrill 7671, which I have not, from the Philip- 
pines. The Carr specimens conform to what there is of a 

20. C. SUPRiiS'TRIGOSUS (Ros.) Copel. 

C suprastrigosus (Ros.) Copel., Genera (1946) 143. 
Dryopteris suprastrigosa Ros., Fedde's Repert. 10 (1912) 335. 

Bamler 87, 4, Sattelberg, distributed as Rosenstock Fil. novog. 
exsicc. n. 28 i, in Herb. Univ. Calif.; Brass 5158, Mafulu, alt. 
1,250 m. Described as having medial pinnae 4 cm long; but 
our specimen, probably Bamler i before distribution, has them 
up to 9 cm long. It differs from C heterocarpus in having 
smaller segments and minutely but densely setulose upper 

21. C. VI^TIGIATUS Copel. Plate 27. 

C vestigiatus Copel., Genera (1946) 143. 

Dryopteris vestigiata Copel., Univ. Calif. Publ. Bot 18 (1942) 220. 

C. rhizomate erecto vel adscendente, basibusque stipitum fas- 
ciculatorum paleis castaneis lanceolatis ca. 4 mm longis sparsis; 
stipite 50 cm alto, 4 mm crasso, f usco sursum glabro vel glab- 
rescente vestigiis nonnullis omnino inconspicuis pinnarum pra- 
edito; lamina 1 m alta vel altiore, 30-35 cm lata, sensim 
acuminata, basi (absque vestigiis pinnarum) truncata, bipinna- 
tifida, rhachi minutissime pubescente; pinnis ultra 40-paribus, 
brevi-pedicellatis, infimis aut deflexis aut horizontalibus, rhachin 
vei^us angustatis, medialibus basi truncatis, usque ad 18 cm 


Copeland: Aspidiaceae of New Guinea 4:4:1 

longis 2 cm latis, caudato-acuminatis, alibi ad alam 2 mm latam 
costae pinnatifida, herbacea, segmentis 3.5-4 mm latis, obtusis, 
superne costis ochroleuco-strigosis venis sparse et decidue albo- 
setosis lamina minutissime decidue setulosa, inferne costis glab- 
rescentibus aliter glabris; venis ca. 9-paribus, infimis anastomo- 
santibus, sequentibus ad marginem supra sinus rotundun attin- 
gentibus ; soris medialibus, indusiis f ugacibus vel nullis, sporan- 
giis nudis. 

Dutch New Guinea : 15 km S.W. of Bernhard Camp, Idenburg 
River, alt. 1,600 m. Brass 12317, "Plentiful in undergrowth of 
rain-forest gullies; fronds somewhat over 2 m long, spreading 
from a slender stock up to 60 cm long." 

Apparently related to C. alatellus and C. heterocarpus; 
nearest to C. ohlongus of the several similar species described 
by Brause. There seems to be hardly a limit to the number 
of similar but distingishable species in this group. 

22. C. OBLONGUS (Brause) Copel., comb. nov. 

Dryopteris ohlonga Brause, Engler's Jahrb. 56 (1920) 109. 

Known only by the type collection, Ledermann 10100, Sepik 
region, alt. 1,000 m. Epiph3rtic; distinguished from C. auste- 
rits by abrupt contraction of base of frond, and more deeply 
cut pinnae. 

23. C. SUPERBUS (Brause) Copel., comb. nov. 

Dryopteris superba Brause, Engler's Jahrb. 56 (1920) 105. 

Known only by the original collections, Ledermann 11738 
and 118^8, Sepik region, alt. 2,070 m. Terrestrial; lamina 125 
cm long; medial pinnae 17 cm long; lower pinnae rather abruptly 
reduced to rudiments. 

24. C. HETEROCARPUS (Blume) Ching 

C. heterocarpus (Blume) Ching, Bull. Fan Bot. 8 (1938) 180. 

Reported, as Dryopteris heterocarpa, Ledermann 11601. The 
only New Guinea specimen I have seen under this name was 

Cyclosonis sp.. King iOi, from Ambasi, apparently unde- 
scribed, is a member of the preceding group, with a few lowest 
pinnae abruptly reduced to rudiments, costae naked beneath, 
and indusia fugacious or none. 

25. C. SUPRASPINIGERUS (Ros.) Copel. 

C supraspinigerus (Ros.) Copel., Genera (1946) 148. 
Dryopteris supraspinigera Ros., Hedwigia 56 (1915) 353. 

Bamler 91 (1913), Sattelberg, alt. 900 m; isotype in Herb. 
Univ. Calif, Not again collected. 

10832 6 

448 The Philippine Journal of Science ^^^^ 

26. C. PARAPSrrSATUS €o9el. 

C paraphysatus CopeL, Genera (1946) 143. 

Dryopteris paraphysata CopeL, Philip. Jour. Sci. 6 C (1911) 74. 

King 806, Papua. With this may probably be identified Brass 
18886 and 18887, Idenburg River, alt. 150 m, in a rocky ravine 
on lower mountain slopes. These specimens show moderate 
dimorphism, and it is the fertile frond which is like the single 
frond of the type. Sterile pinnae reach 23 cm in length and 
3.5 cm in width, cut one-third of the way to the costa. The 
rhizome is stout, creeping, clothed at the apex with fuscous 
pubescent paleae. 

27. C. PARIPINNATUS Copel. Plate 28. 

C. paripinnatus CopeL, Genera (1946) 143. 

Dryopteris paripinnata CopeL, Univ. Calif. PubL Bot. 18 (1942) 220, 

C. rhizomate erecto, apice basibusque stipitum haud conges- 
torum paleis fuscis membranaceis integris glabris lanceolatis 
vix 1 cm longis vestitis ; stipite usque ad 1 m alto, f usco, glab- 
rescente, sursum rudimentis nonnullis remotis pinnarum pra- 
edito; lamina ca. 80 cm alta, basi truncata, pinnis oppositis, 
sursum in segmenta apicis acuminati sensim transeuntibus, 
infimis expansis et medialibus ca. 20 cm longis, 2-2.5 cm latis, 
in apices acuminates integros subf alcatos protractis, subsessili- 
bus basibus pauUo angustatis, subcoriaceis, costas versus ultra 
medium pinnatifiidis, segmentis rotundatis ca. 5 mm latis ; rhachi 
fulvo-setosa, costis utrinque leviter setosis, costulis superne 
nudis inf erne venisque breviter sparse setosis, lamina superne 
glabra inf erne minute sparse setulosa; venis usque ad 9-paribus, 
infimis anastomosantibus, sequentibus 1-paribus conniventibus 
et plerisque anastomosantibus, ceteris secus marginem curvis; 
soris inframedialibus, indusiis glabris, sporangiis non spinuli- 

Dutch New Guinea: 15 km S.W. of Bemhard Camp, Idenburg 
River, alt. 1,700 m. Brass 12A85, type, "a large clump-fern 
common in rain-forest gullies;'' No. 12156, ibidem. 

Remarkable for its height and its rather remote, strictly 
opposite pinnae. It differs from C. appendicuMus in much 
firmer texture, pinnae contracted at base, and in other details. 
Other Cychsorus species with opposite pinnae are C. hiauritus 
in India, C. malodorus in the Solomons, C. pennigerus in New 
Zealand, none evidently related to C paripinnatus. Affinity to 
the group of C crassifolius is likewise not apparent. 

Carr 10202 represents a relataed species, with few, relatively 
broad pinnae less deeply lobed, its veins sparsely setose above. 

'^'^' ^ Copehnd: Aspidiaceae of New Guinea 449 

28. C. DELTIPTEBUS Copel. Plate 29. 

0. deltipterus Copel., Genera (1946) 142. 

Dryopteris deltiptera CopeL, Univ. Calif. Bot. 18 (1942) 220. 

C. paripinnato affiinis, pinnis basi dilatatis et facie inferiore 
glabra distinctus ; rhizomate adscendente, apice basibusque stipi- 
tum confertorum paleis fuscis membranaceis nudis integris ca. 
1 cm longis vestitis; stipite ca. 60 cm alto, brunneo, sursum in 
sulca strigoso alibi glabro, rudimentis nomiullis pinnarum pra- 
edito; lamina usque ad 1 m longa, apice breviter acuminate 
inciso deltoideo basi truncate; pinnis oppositis, remotis, infe- 
rioribus sterilibus 15 cm longis acutis basi 3.5 cm latis truncatis 
sessilibus, subcoriaceis, J— ^ ad costas incisis, segmentis rotun- 
datis integris; rhachi costisque superne strigosis, inferne cum 
facie tota glabris; venis ca. 14-paribus, quorum ca. 6-paribus 
anastomos-antibus, liberis in marginem angustum cartilagineum 
curvis ; f ronde fertile modo contracta, pinnis prof undius incisis, 
venis ca. 10-paribus fere omnibus soriferis; soris inframedia- 
libus, indusiis persistentibus glabris. 

Dutch New Guinea: Bele River, alt. 2,200 m, Brass 11260, 
type in Gray Herbarium, "large ground fern, common in flood- 
plain forest." 

A relative of C. paripinnatus, most conspicuously distinguished 
by the form of the pinnae, uniformly narrowed from the broad 
truncate base to the apex. 

29. C. SUBAPPENDICULATUS Copel. Plate 30. 

C. subappendiculatus CopeL, Genera (1946) 143. 
Dryopteris subappendiculata CopeL, Univ. Calif. PubL Bot. 18 (1942) 

C. rhizomate ignoto ; stipite ad vestigia infima pinnarum 10 cm, 
ad pinnas normales 35 cm alto, basi nigro paleis paucis brunneis 
6 mm longis praedito, alibi rhachique fuscis setis obscuris 
brevibus vestitis ; lamina 90 cm longa, 30 cm lata, subdimorpha, 
basi truncata, segmento apicale 15 cm longo integro-acuminato 
deorsum inciso-lobato, alibi pinnata; pinnis ca. 16-paribus 
normalibus, oppositis, remotis, inferioribus 18 cm longis, 2.5-3 
cm altis, acuminatis, basi truncatis vix dilatatis, sterilibus inciso- 
crenatis lobis 5 mm latis 1.5 mm longis truncatis, papyraceis, 
costis superne appresso-strigosis, inferne glabris, costulis venis 
et lamina glabris ; venis 4-6-paribus plerisque anastomosantibus ; 
pinnis inftmis 6-paribus abrupte in auriculas triangulari-hastatas 
1 cm longas reductis; pinnis fertilibus lobatis, lobis 5 mm latis 
2-3 mm longis f alcato-truncatis ; soris medialibus vel infra- 

450 The Philippine Journal of Science i^^^ 

medialibus, indusio inconspicuo brunneo setis paucis ornato, 
sporangiis nudis. 

Dutch New Guinea: 15 km S.W. of Bernhard Camp, Idenburg 
River, alt. 1,700 m, Brass 122^7; "Several clumps in under- 
growth of rain-forest gulley; fronds dimorphous, ib 1 m long." 

Near C. deltipteruSy distinguished by longer, narrower pinnae, 
with parallel and less incised margins, the rudimentary pinnae 
below the base of the normally developed frond larger and 

29a. C. APPENDICULATUS (Blume) CopeL 

0. appendiculatus (Blume) CopeL, Genera (1946) 142. 

Reported from New Guinea. I have seen no specimen, and 
suspect wrong identification. 

30. €. SOGERENSIS (Gepp) Copel. 

C sogeremis (Gepp) Copel., Genera (1946) 143. 
Dryopteris soger ensis Gepp, Jour. Bot. 61 (1923) Suppl. 61. 

Described from collections by Forbes at Sogere, Papua, alt. 
1,700-2,500 feet; pinnae 15-19 cm long, 1.5-2 cm wide; not 
well distinguished by description from C. paraphysatus. But, 
identified as this species, comes Carr H667, Boridi, alt. 3,500 
feet, pinnae 30 cm long, 3-3.5 cm wide, distinguishable in many 
ways. Identical with this specimen is King ill, from Mamba, 


31. C. EXTENSUS (Blumc) Ching 

C. extensm (Blume) Ching, Bull. Fan Bot. 8 (1938) 182. 
Aspidium extensum Blume, Enum. (1828) 156. 

Schlechter 163i9, alt. 180 m. Our specimen of this collection 
might not be identified, but bears this name, and it may well 
be correct. 

Supposed to range from New Caledonia to Africa. 

31a. CTCLOSORUS sp. 

Brass 5197, in Herb. N. Y. Bot. Garden as Dryopteris hispi- 
dvliformis (No. 16), has the lower pinnae abruptly reduced, 
1 or 2 anastomosing veinlets above the lowest pair, and lamina 
setulose beneath. It is near C tadavensis of Mindanao, but 
not identical. 

32. C. MEGAPHITLLOIDES (Ros.) Copel. 

C. megaphyUoides (Eos.) Copel., Grenera (1946) 143. 
Dryopteris megaphylloides Ros., Fedde's Repert. 12 (1913) 174. 

Keysser 120, Sattelberg, alt. 800-1,000 m. Our specimen is 
labelled as Bamler 120^ almost surely a mistake. Also (var. 

''"^'^ Copeland: Aspidiaceae of New Guinea 451 

glabrescens Eos.) Rosenstock Fil. novog, exsicc, n, 233. King 
487, mountains behind Medan, is approximately this species, 
more setose than the type, and with broader lobes. 

32a. CYCLOSORUS sp. 

Carr H202, Boridi, alt. 5,000 feet, received as D. mega- 
phylMdes and conforming reasonably to the description, is 
really no near relative; it is probably undescribed. 

33. C. ATASRIPn (Ros.) Copel. 

C. Atasripii (Ros.) Copel., Genera (1946) 142. 
Dryopteris Atasripii Ros., Meded. Rijks Herb. No. 31 (1917) 6; 
C. Chr., Brittonia 2 (1937) 298.* 

Brass 13844, Idenburg River, alt. 85 m, frequent on lower 
forested slopes; Bamler 51 (1912) Logaueng, alt. 400 m, det. 
Rosenstock; Brass 5153, det. Christensen, and the subject of 
the first adequate description of the species. Here belongs also 
Schlechter 16855, alt. 500 m, received as D, sagittifolia. It 
thus occurs at minor and middle altitudes throughout the island. 


Dryopteris tamiensis Brause, Engler's Jahrb. 49 (1912) 23. 

Known only by the sterile type, Schultze (33) 86, from the 
interior of Kaiser-Wilhelmsland, ait. 1,600 m. 

35. C. ARFAKIANUS (Baker) Copel. 

C. arfakianus (Baker) Copel., Genera (1946) 142. 
Dryopteris arfakiana (Baker) C. Chr., Index (1905) 253; Dansk Bot. 
Arkiv 9 No. 3 (1937) 50. 

King 483, and (?) 485, from mountains behind Medan; Carr 
12021, Koitake, Papua, alt. 1,500 feet. So far as my specimen 
shows, Carr 15531, Isuarava, alt. 4,500 feet, is the same species. 
Here also Schlechter 17743 and 17746, Kani Mountains, alt. 
1,000 m, and Bamler, Rosenstock Fil novog. exsicc. n. 231, 
Sattelberg, alt. 900 m, all received as Dryopteris arhuscula, but 
with little resemblance to the fern properly so called. Christen- 
sen regards D. sepikensis as a small form of this species. King 
483 has an erect caudex 20 cm tall; the paleae are puberulent. 


36. C. GREGARIUS Copel. Plate 31. 

C gregarius Copel., Jour. Arnold Arb. 24 (1943) 440. 

Known only by the type collection, Brass 6759, from the upper 
Fly River, alt. 80 m. 

452 The Philippine Jounml of Science ^^^^ 

36A. C. CYATHEOIDl^ (Kaalfuss) FarweU 

C. cyathemdes (Katilfuss) Farwell, Amer. Midland Nat. 12 (1931) 258. 

Reported from New Guinea, but probably in error; so far 
as I know it, this species is endemic in Hawaii. 

37. C. UNITUS (L.) Ching 

C, unitus (L.) Ching, Bull. Fan Bot. 8 (1938) 196. 

Brass 11587, 13250, probably 2,200 and 850 m; the former 
altitude is notable for this species, but the location was grass- 
land, formerly cultivated. 

Common, to Polynesia and East African islands. 

38. C. ARmUS (Don) Ching 

C. aridus (Don) Ching, Bull. Fan Bot. 8 (1938) 194. 

Papua, King 172, 40S; Brass U07; Kaiser-Wilhelmsland, 
Schlechter 16777. 
To India and Fiji. 

39. C. ARCHBOLDH (C. Chr.) Copel. 

C. Arehholdii (C. Chr.) Copel., Genera (1946) 142. 
Dryopteris Arehholdii C. Chr., Brittonia 2 (1937) 297. 

Papua: Mt. Tafa, alt. 2,400 m, Brass 4875. Collected only 


DryopteHs pUososquamata v.A.v.R., Bull. Agric. Ind. Ind. N^rl. Ill 
(1908) 4. 

Described from a cultivated plant in the Buitenzorg Garden, 
originally from Skroe, New Guinea. 

41. C. MEGAPHYLLUS (Mett.) Ching 

C. megaphyllus (Mett.) Ching, Bull. Fan Bot. 8 (1938) 225. 

Unknown to me in New Guinea. Cited are Ledermann 7553, 
8316, and, as var. abbreviata Ros., Keysser 262. 
To the Philippines and Africa. "" 

42. C. TRUNCAT0S (Poiret) FarweU 

a truncatm (Poiret) Farwell, Am. Midland Nat. 12 (1931) 259. 

Brass 12221, 12273, alt. 1,750 m; Bamler, Rosenstock Fil. 
novog, exsicc. n. 235; Brass 5m, Papua, alt. 1,250 m. 
To India. 

43. C. RIGmUS (Ridley) Copel., comb. nov. 

Goniopteris rigida Bidley, Trans. Linn. Soc, Bot. II 9 (1916) 258. 
Cyclosorm Wollastonii (v.A.v.R.) Copel., (»enera (1946) 143. 

The original description of this species was very inadequate; 
However, it has a type, of which Mr. Alston has been good 

^^' ^ Copeland: Aspidiaceae of New Guinea 453 

enough to send me a photograph. Brass 10287, 10826, Lake 
Habbema, alt. 2,800 m, resemble it just enough so that I do not 
care to give them a new name. Because the identification is 
doubtful, I will not correct and amplify the description, more 
than is done in the key to the species. 

44. C. MULTIAURICULATUS Copel. Plate 32. 

C. multiauriculatus CopeL, Genera (1946) 143. 
Dryopteris multiauriculata CopeL, Uinv. Calif. Bull. Bot. 18 (1942) 

C. caudice stricte erecto, stipitibus et radicibus exclusis 1 cm 
diametro, apice stipitibusque paleis rufo-ferrugineis ca. 7 mm 
longis lanceolatis pubescentibus vestitis; stipitibus 5-10 cm 
longis, caudici appressis; frondibus subdimorphis, acuminatis, 
deorsum longe gradatim usque ad apicem caudicis angustatis; 
fronde sterile 80-100 cm alta, 20 cm lata, rhachi fusco-setosa; 
pinnis permultis, alternantibus, sessilibus, basi truncatis, acutis, 
medialibus 10 cm longis, basi leviter dilatatis 1.5 cm latis, 
integris vel obscure crenulatis, firme papyraceis, costis supeme 
sparse inferne densius setulosis, alibe setulosis setulis caducis; 
venis 4- 5-paribus, plerisque anastomosantibus; pinnis superio- 
ribus in segmenta apicis sensim transeuntibus ; fronde fertile 
pauUo minore, venis 3- 4-paribus omnibus sorif eris, soris media- 
libus, primo orbicularibus deinde nonnulis elongatis, infimis 
saepe more Meniscii confluentibus, tum demum f aciem inf eriorem 
omnino tegentibus, indusiis invisis, sporangiis nudis. 

Dutch New Guinea: 6 km SW. of Bernhard Camp, Idenburg 
River, alt. 1,150 m, Brass 12850, type, "common ground fern 
in rain-forest gullies; fronds dimorphous, spreading from an 
erect stock up to 40 cm in length;" also. No. 1305 i, alt. 850 m. 

Similar, and probably related, to C. hastato-pinnatm, No. 
62, which has a symmetrical entire terminal leaflet, and erecto- 
patent lateral pinnae pronged on the upper side, with the surface 
probably naked from the start; they are alike in the gradual 
shortening of the crowded lower pinnae, the lowest broadly 

45. C. PROTECTUS Copel. Plate 33. 

C. protectua Copel, Genera (1946) 143. 

Dryopteris protecta CopeL, Univ. Calif. Publ. Bot. 18 (1942) 221. 

C. caudice erecto, 10-12 mm diametro, apice dense et stipitibus 
sparsius paleis f uscis 7-10 mm longis late lanceolatis minutissime 
decidue puberulentibus vestitis; stipitibus ad vestigia infima 
pinnarum 5-10 cm altis; lamina 70-SO (basi contracta exclusa 
ca. 50 cm) alta, 14 cm lata, apice pinnatifida acuminata, basi 

454 The Philippine Journal of Science ^^^^ 

sensim vel subabrupte contracta, pinnis infimis remotis ad 
vestigia vix manifesta reductis, rhachi f errugineo-setulosa ; 
pinnis medialibus altemantibus, subsessilibus, 7.5 cm longis, 
13 mm latis, acutis, basi truncatis, one-fourth to one-third ad 
costam incisis, segmentis 2.5 mm latis, subcoriaceis, costis 
breviter strigosis, costulis venisque sparse setulosis, lamina 
glabra; venulis ca. 5-paribus, biparibus plerumque anastomo- 
santibus; soris inframedialibus (hand costularibus), indusiis 
persistentibus setulosis. 

Dutch New Guinea: 9 km N.-E. of Lake Habbema, alt. 2,800 
m, Brass 109 SS, type; "Abundant in thin secondary forest on 
an old landslip; leaves spreading from a semi-erect slender 
stem 40-50 cm high;" No. 112A6, Bele Kiver, alt. 2,300 m; 
No. 12Jf70, on an open rock-slide, alt. 2,150 m, fronds only 
45 cm long and hardly 10 cm wide. 

46. C. AMBOINENSIS (WUld.) Copel. 

C. amboinensis (Willd.) Copel., Genera (1946) 142. 

C. suhpubescens (Blume) Ching, Bull. Fan Bot. 8 (1938) 211. 

Both of these species, as well as Dryopteris mollis which has 
been used as a name of both, have been reported from New 
Guinea. Carr 11635, from Veiya, Papua, comes as C. suhpubes- 

C. dentatus (Forsk) Ching, which, broadly construed, can 
include all of these, is then pantropic. 

47. C. TERRESTBIS Copel. Plate 34. 

C. terrestris Copel., Genera (1946) 143. 

Dryopteris terrestris Copel., Univ. Calif. Publ, Bot. 18 (1942) 221. 

C. rhizomate brevi-repente, basibusque stipitum paleis atro- 
f uscis ca. 4 mm longis microscopice setulif eris vestitis ; stipitibus 
approximatis, usque ad vestigia infima pinnarum 10-15 cm, 
usque ad pinnas normales 30 cm altis, rhachibusque minute 
appresso-setosis ; lamina (basi contracta exclusa) ca. 45 cm 
longa, 15 cm lata, apice 15 cm longo pinnatifido acuminato; 
pinnis infimis paucis remotis ad vestigia reductis, infimis nor- 
malibus deflexis, 5-6 cm longis, media longitudine quam basi 
latioribus; medialibus 7-8 cm longis 2 cm latis, altemantibus, 
sessilibus, basi recte truncatis, abrupte falcato-acuminatis et 
hie integris, alibi serratis vel inciso-serratis, papyraceis, micros- 
copice sparse setulosis oculo nudo glabris ; venulis 5-paribus, 
plerisque anastomosantibus et soriferis; soris medialibus, in- 
dusiis et sporangiis setuliferis. 

Dutch New Guinea: 2 km S.-W. of Bemhard Camp, Idenburg 
River, alt. 700 m, Brass 13660; "Rain forest; common ground 

"^■^'^ Copeland: Aspidiaceae of New Guinea 455 

fern on steep slopes;'' The illustration, Pfate 35, shows the 
peculiar form of the pinnae, better than is possible with words. 

48. C. CONFERTUS (Brause) Copel. 

C. confertus (Brause) CopeL, Genera (1946) 142. 

Dryopteris conferta Brause, Engler's Jahrb. 49 (1912) 22, /, 1 F. 

Kani-Gebirge, alt. 700 m, Schlechter 178A6, isotype in Herb. 
Univ. Calif. Brass 38^9, Dieni, Central Division, Papua, alt. 
500 m, is superficially similar, but is long-stipitate, and has 
minute indusia and naked sporangia. Brause's figure of the 
whole frond gives little idea of its appearance. 


49. C. DICHBOTRICH17S Copel. 

C dichrotrichus Copel., Genera (1946) 142. 

Dryopteris dichrotricha Copel., Philip. Jour. Sci. 6 C (1911) 74. 

Papua, King 2H, 219; otherwise unknown. This species 
and D. dichrotrichoides v.A.v.R., a Philippine species, are 
reversed by Christensen, Suppl. Ill 84. 

50. C. mSPIDULUS (Dec.) Copel. 

C hispidulus (Dec.) Copel., Genera (1946) 142. 

Aspidium hispidulum Decaisne, Nouv. Ann. Mus. Paris 3 (1834) 346. 

Dryopteris angusta Copel., Philip. Jour. Sci. 9 C (1914) 3. 

King 408, type of D. angitsta, from Papua Schlechter 16805, 
alt. 300 m; Bamler, Rosenstock Fil. novog. exsicc. n. 232, alt. 
800 m. Dryopteris perpilifera V.A.V.R., No. 15 of this list may 
be another synonym. 

Malaya; Samoa. 

51. C. ALBOSETOSUS Copel. Plate 35. 

C, albosetosus Copel., Genera (1946) 142. 

Dryopteris albosetosa Copel., Univ. Calif, Publ. Bot. 18 (1942) 221. 

C. rhizomate brevi-repente, 3 mm crasso, basibusque stipitum 
paleis atrocastaneis ovatis minute ciliatis vestitis; stipitibus 
approximatis, usque ad vestigia infima pinnarum ca. 5 cm (ad 
laminam expansam 10-15 cm) altis, sursum rhachibusque bre- 
viter cinereo-setosis et pulverulentibus ; lamina (pinnis infimis 
ca. 6-paribus subabrupte reductis exclusis) 20-25 cm longa, 7 cm 
lata, sursum in apicem acuminatum pinnatifidum 7-9 cm longum 
sensim angustata; pinnis infimis normalibus paullo abbreviatis 
vix deflexis, medialibus altemantibus subsessilibus, 4 cm longis, 
1 cm latis, obtusis vel subacutis, basi truncatis, inciso-dentatis 
dentibus 3 mm latis margine deflexis, subcoriaceis, superne 
ubique albosetosis; venulis 5- 6-paribus quarum 3-paribus an- 
astomosantibus; soris medialibus, indusiis et sporangiis setosis. 

456 The Philippine Journal of Science ^^^^ 

Dutch New Guinea: Bele River, alt. 2,200 m, Brass 1US5; 
"Abundant in small clumps on dry face of a limestone wall in 
forest shade." 

Related to C. confertus, but pinnae smaller, more incised and 
more setose. 

52. C. STBIGOSISSIMUS Copel. Plate 36. 

C. strigosissimus Copel., Genera (1946) 143. 

Dryopteris strigosissima Cope}., Univ. Calif. Publ. Bot. 18 (1942) 221. 

C. rhizomate erecto, breve, stipitibusque paleis atrofuscis 
ovatis acuminatis minute pubescentibus usque ad 8 mm longis 
vestitis; stipitibus fasciculatis, 2-4 cm longis; lamina 40 cm 
longa, 10 cm lata, apice acuminato 2 cm et pinnatifido 4 cm 
longo, basi angustata et deinde in crus 10 cm longum contracta, 
pinnis hie contiguis vel imbricatis sursum 1 cm longis triangulari- 
hastatis deorsum sensim ad vestigia diminutis, rhachi dense 
fulvo-tomentosa; pinnis medialibus altemantibus sessilibus, acu- 
minatis, basi pauUo dilatatis 8-10 mm latis fere contiguis, 
dentatis dentibus 1 mm longis 2 nmi latis, papyraceis, facie 
superiore ubique densissime breviter pallide setosa; venulis ca. 
5-paribus, 3-paribus anastomosantibus ; soris medialibus, indusiis 
magnis persistentibus dense setosis, sporangiis glabris. 

Dutch New Guinea: Bele River, alt. 2,200 m., Brass 1U36. 
"A few tufted plants on dry face of a limestone cliff in forest 

It will be observed that this and the preceding species were 
collected together and look much alike. Two superficially similar 
Mexican species grow on shaded limestone cliffs. 

53. C. RIPAKIUS Copel. Plate 37. 

C riparius CopeL, Genera (1946) 143. 

Dryopteris riparla CopeL, Univ. Calif. Publ. Bot. 18 (1942) 221. 

C. rhizomate adscendente, 4 mm crasso, basibusque stipitum 
paleis paucis fuscis ovatis subglabris 2 mm longis vestitis; 
stipitibus confertis, usque ad pinnas infimas remotas vestigiales 
5-12 cm altis, pube decidua et paleolis paucis caducis praeditis ; 
lamina 40 cm longa, 7-9 cm lata, apice acuminato pinnatifido 
breve, basi sensim vel abrupte contracta pinnis reductis paucis, 
rhachi supeme strigosa, inf erne setulosa vetustate glabrescente ; 
pinnis medialibus altemantibus subsessilibus, 5.5 cm longis, vix 
1 cm latis acutis basi cuneatis interdum subauriculatis, 
Y^ ad costam inciso-serratis dentibus 3 mm latis 1.5 mm 
longis, subcoriaceis, costis supeme strigosis, aliter glabris; 
venulis biparibus anastomosantibus, 2 3-paribus liberis; soris 
medialibus, indusiis persistentibus sporangiisque glabris. 

'^'^'^ Copeland: Aspidiaceae of New Guinea 467 

Dutch New Guinea: 4 km S.W. of Bernhard Camp, Idenburg 
River, alt. 800 m, Brass 130^8. "Very abundant on steep, 
shaded, flood-swept bank of river/' 

There is some resemblance to the Philippine C. philippinensis, 
but this may be due wholly to the conmion habitat. Typically 
flood-swept species look alike, the world over. 

54. C. IMPONENS (Ces.) Copcl. 

C. imponens (Ces.) CopeL, Genera (1946) 142. 

Dryopteris imponens (Cesati) C. Chr.; see Dansk Bot. Arkiv 9 
No. 3 (1937) 50. 

D. armata Ros. Hedwigia 56 (1915) 351. 

D, muricata Brause, Engler's Jahrb. 56 (1920) 106. 

Brass 12U8, 18269, 18728, alt. 800-1,200 m; Carr 15U0, 
Isuarava, alt. 4,500 feet; Bamler 111, Sattelberg, alt. 700-800 
m, distributed as Rosenstock Fit novog. exsicc. n. 2^2, both in 
Herb. Univ. Calif. King 455 is near this species, not identical. 

D. muricata is reduced by description. Brause distinguished 
it from D. armata: 

1. By the basal paleae. In this respect, Bra^s 12H8, his 
only specimen with stipe base, fits Brause's description. Our 
stipe base of Bamler 111 is that of a small frond, certainly of 
the same species. 

2. By the spacing of the pinnae, described by Brause as 
"subcontiguis." They are contiguous on the upper part of the 
frond of Bamler 111, but are rather remote toward the base 
of specimens showing the base; Brass 12948 shows both base 
and apex. Brause's specimen was evidently incomplete, — "Folia 
usque ad 2 m (e scida) longa." Brass' note on No. 13269 shows 
a total height of 4 meters. 

3. By elongate sori, especially the lower ones. This is more 
or less evident on all specimens. 


55. C. INTERRUPTUS (Willd.) Ching 

C. interruptus (Willd.) Ching, Bull. Fan Bot. 8 (1938) 184. 
Aspidium pteroides Sw., non Poly podium pteroides Ketz. 

Carr 11123, 11722, Papua, at sea level. 
To India and China; and reported from Queensland and 

56. C. mVISUS (Forster) Copel. 

C invistis (Forster) Copel., Genera (1946) 142. 
Polypodium invisum Forster, Prod. (1786) 81. 

Brass 5538, Mafulu, Papua, alt. 1,250 m; previously reported. 
Polynesia, Philippines. 

458 The Philippine Journal of Science i^^^ 

57. C. GONGYLODES (Schkuhr) Link 

C. gongylodes (Schkuhr) Link, Hort. Berol. II (1833) 128. 

Brass H076, alt. 50 m. "Massed in floating grass of deep, 
open marshes ; common.'' It should be common in wet meadows, 
but seems not to have been reported before in New Guinea. 

All warm lands; New Zealand. 

58. C. GLANDULOSUS (Bliime) CMng 

C. glandulosus (Blume) Ching, Bull. Fan Bot. 8 (1938) 227. 

Reported from New Guinea but I have seen no specimen. 
Malaya; Philippines; Solomon Islands. 

59. C. CUSPIDATUS (Blume) Copel. 

C cuspidatus (Blume) Copel., Genera (1946) 142. 
Meniscium cuspidatum Blume, Enum. (1828) 114.^ 

I have seen no specimen; Brause cites Ledermann 974S as 
Dryopteris urophylla, var. ctispidata. 
Malaya; Philippines. 

60. C. imOPHYLLUS (WaU.) Copel. 

C. urophyllus (Wall.) Copel., Genera (1946) 143. 

King 258, Papua; Bamler, Rosenstock FiL novog. exsicc. n. 
17 Jf, Sattelberg, alt. 900 m. There is also var. peraspera 
v.A.v.R., Lam 677, not seen. 

Asia to Australia and Fiji. 


Dryopteris rufo-pilosa Brause, Engler's Jahrb. 56 (1920) 106. 
Goniopteris rudis Ridley. Trans. Linn, Soc. Bot. II 9 (1916) 259. 

A photograph of the type, kindly sent by Mr. Alston, shows 
the pinnae to be alternate, the veinlets up to 11 pairs, the 
soti medial, — all contrary to the description. I do not see why 
it is not C. urophyllus. 

61. C. MICANS (Brause) Copel. 

C micans (Brause) Copel., Genera (1946) 143. 
Dryopteris micans Branse, Engler's Jahrb. 56 (1920) 98. 

Brass 12900, 18392, alt. 850 and 1,150 m. Described from 
Ledermann 12468 and 11278. Endemic. 

62. C. HASTATO-PINNATUS (Brau^) Copel. 

C. hastatO'pinnatus (Brause) Copel., Genera (1946) 142. 
Dryopteris kastato-pinnata Brause, Engler's Jahrb. 56 (1920) 112. 

Brass 13419, alt. 850 m, "scattered in the stony bed of a 
rain-forest stream." Described from Ledermann 8237, 70U7. 

'^^'^ Copeland: Aspidiaceae of New Guinea 459 

Suggestive of C aqiuitilis and C. riparia, in adaptation to the 
common habitat. 

63. C. MOROBENSIS Copel. Plate 38. 

C. morobensis Copel., Genera (1946) 143. 

Dryopteris morobemis Copel., Univ. Calif. Publ. Bot. 18 (1942) 221. 

Pseudomeniscium trunco arboreo 150 cm alto (teste lectore) ; 
stipite 20 cm alto 6 mm crasso, f usco, basi paleis f uscis appressis 
integris membranaceis 1 cm longis vestito, alibi glabrescente 
asperulo ; lamina ultrametrale subdimorpha, pinnata, basi sensim 
longe angustata, rhachi glabra ad basin pinnae quaeque aero- 
phoro praedita; frondis sterilis pinna apicale obscure crenata 
aliter ceteris conforme, pinnis inferioribus gradatim reductis 
et remotis, infimis vix 5 mm longis et latis; pinnis medialibus 
permultis alternantibus, erecto-patentibus, 20-25 cm longis, 
1.5-2 cm latis, sensim acuminatis, subsessilibus, basi acroscopica 
cuneata basiscopica rotundata, integris vel sinuatis, subcoriaceis, 
glabris; venulis 3- 4-paribus oblique anastomosantibus, 1- 
2-paribus brevibus liberis; frondis fertilis partibus omnibus 
minoribus, pinnis ca. 12 cm longis 8 mm latis, crenatis, venulis 
ubique soriferis, soris nudis confluentibus paginam inferiorem 
dense tegentibus. 

New Guinea: Morobe, Finongan, alt. 1,400 m, Clemens s. n,, 
*'m mossy mountain bush; trunk 5-6 feet." 

This seems much like C. dimorphus (No. 6), but the descrip- 
tion of that species is explicit, "nervis • . . infimis anastomos- 
antibus," and "das unterste Paar anastomosierend." The sori 
are not merely crowded as the sporangia mature; they are some- 
what indefinite, more or less elongate along the veinlets, and 
with border-sporangia on the "parenchyma." 

64. C. TRIPHYLLUS (Sw.) Copel. 

C. triphyllus (Sw.) Copel., Genera (1946) 143. 
Meniscium triphyllum Swartz, Schrader's Journal (1801) 16. 

Papua: King 322; Carr 11983. 
Queensland to India. 

65. C. PENTAPHYLLUS (JRos,) Copel. 

C pentaphyllus (Ros.) Copel., Genera (1946) 143. 
Dryopteris pentaphylla Ros., Fedde's Repert. 12 (1913) 529. 

Known only by the type collection, Keysser 186 (1913) 
Hinterland des Sattelbergs, 1,400-1,500 m; isotype in Herb. 
Univ. Calif. 

460 The Philippine Journal of Science ^^^^ 

€#. €L CANESCSNS (Blmne) Copel. 

C eanescens (Bltime) Copel., Genera (1946) 142. 
Dryopteris eanescens var. novo-guineenssis Brause, Engler's Jahrb. 
40 (1912) 22. 

Brass 12210, alt. 1,750 m. ''Bank of a forest stream; common 
on rocks and on ground." The varietal type is Schlechter 188H, 
Bismarck Mountains; isotype in Herb. Univ. Calif. 

The variety is endemic, and would better be regarded as a 
specie, as I have treated nearly related Philippine members 
of the group. It is distinguished from the supposedly typical 

C. eanescens by some degree of dimorphism and by having only 
the lowest veinlets anastomosing. 

67. C. LANCEOLUS (Christ) Copel. 

C lanceolus (Christ) Copel., Genera (1946) 142. 

Dryopteris lanceola (Christ) Copel., Philip. Jour. Sci. 56 (1935) 102. 

Keysser II 85 (1912), Bolan, alt. 2,400-3,000 m, received as 

D. eanescens var. nephrodiiformis Christ; fertile frond only, 
exactly like Philippine fertile fronds. 


68. C. BECCABIANUS (Ces.) Copel. 

C beccarianus (Ces.) Copel., Genera (1946) 142. 

Meniscium beccarianum Cesati, Rend. Ac. Napoli 16 (1877) 27. 

Dryopteris cesatiana C. Chr., Index (1905) 257. 

D. oblanceolata Copel., Philip. Jour. Sci. 9 C (1914) 3. 

D. eanescens var. incana Ros., in herb. 

Brass 12199, 13050, alt. 1,750 and 850 m. 
Throughout New Guinea, evidently common. 


Dryopteris mamberamensis V.A.V.R., Bull. J, B. Buitenzorg II No. 24 
(1917) 3. 

A. PBOLIFEBA (Retz.) Coiie]. 

A. proUfera (Retz.) Copel., Genera (1946) 144, 
Hemuinitis prolifera Retz., Obs. VI (1791) 88. 
Dryopteris prolifera C. Chr., Index 286, and other recent writers. 

Brass 8875, 13936, swampy places near sea level; Lam 1105, 
llil, same region and habitat; King 128, Carr 12285, Papua. 
Polynesia to Africa. 

S. POLYCABPA (Blunie) Copel. 

5. polycarpa (Blume) Copel., Univ. Calif. Publ. Bot. 16 (1929) 60. 

'^''»* Copeland: Aspidiaceae of New Guinea 461 

King 159, 410, 413; Brass 544, all from Papua; Bander, 
Rosenstock FiL novog, exsicc. n. 81, Logaueng, alt. 300 m. 
Larger than known elsewhere, but not otherwise distinguishable. 

Malaya. The generic range includes the Solomon Islands. 

27A. CYSTOPTERIS Bernhardi 
C. TENUISECTA (Blume) Mett. 

C tenuisecta (Blume) Mett. 

Ledermann 858 is reported as this species, but it is misplaced 
in Cystopterls. 
To Formosa and India. 

28. AXHYRIUM Roth 

Key to the species 
Veins free. 
Frond tripinnatifid or more dissected. 
Small ferns, mostly under 20 cm tall. 

Paleae and lamina brownish 1. A, minutum 

Entire plant blackish 15. A, squamuUgerum 

Size moderate, — ^up to 60 cm tall. 
Axes light-brown. 

Pinnae obtuse to acute 2. A, setiferum 

Pinnae acuminate S. A, Myer-Dreesii 

Ax^ dark-brown..... 4. A» scotinum 

Large ferns. 
Fully tripinnate. 
Secondary pinnules incised. 

Sori elongate 7. A. Huttoni 

Sori short, basal 9. A, decompositum 

Secondary pinnules entire. 

Rachises furfuraceous 8. A, latilobum 

Rachises naked except at nodes 6. A. pedicellatum 

Sori short. 

Segments 4 mm wide 10. A, woodwardioides 

Segments 2-3 mm wide 11. A. sylvatictim 

Sori elongate. 
Indusium lacerate. 

Veins nearly all simple 12. A. Blumei 

(Forked veins not rare) 
Forked veins not rare. 

Pinnules 25 mm wide 13. P.* Nymani 

Pinnules 15 mm wide 14. D, Schraderi 

Indusium entire. 

Segments 12 by 3.6 mm 15. D, Schlechteri 

Segments 10 by 5 mm 16. D, Schultzei 

Frond bipinnate, not nearly tripinnate. 
Lamina thin. 
Sori short 2. A. setiferum 

* This key "D" stands for Diplasium, 


The Philippine Journal of Science 


Sori elongate ,. 5. A, morpbense 

Lamina firm. 

Small fern, mostly under 20 cm tall 17. A. squamuligerum 

Much larger. 

Pinnules cut two-thirds to costa 18. P. opadfoUum 

Pinnules cut one-third or one-half to costa. 
Pinnules up to 6 cm long. 

Lamina over 50 cm long 19. A. nitena 

Lamina under 30 cm long 5. A. morobense 

Pinnules commonly 12 cm long 20. A. davaoense 

Pinnules more shallowly toothed. 

Pinnules over 10 cm long 21. A. maximum 

Pinnules 3-5 cm long 22. A. cyatheifolium 

Pinnae pinnate at base, elsewhere subpinnate. 

Paleae of stipe elongate, short-ciliate 23. A. uncidens 

Most paleae reduced to tufts of hairs 24. A. KeysseH 

Pinnae deeply pinnatifid. 
Stellate paleae present. 

Indusium linear 25. A. protensum 

Indusium wanting 26. D, asterotkrix 

Stellate paleae absent. 

Stipe and rachis dark. 

Pinnae about 2.5 cm wide 27. A. sorsogonense 

Pinnae 6 cm wide 28. A. Archboldii 

Axes stramineous or greenish 29. A. japonicum 

Pinnae icised or lobed. 
Pinnae subcordate. 

Lacerate squamules present 30. A. Weinlandii 

Lacerate squamules wanting 31. A. serrato-crenatum 

Pinnae truncate or cuneate at base. 

Sori elongate 32. A. bulbiferum 

Sori horse-shoe-shaped 33. A. Ledeirmanni 

Pinnae serrate 34. A. pallidum 

Pinnae crenate-sinuate or entire. 
Pinnae about 1 cm wide. 

Stipe and rachis stramineous 35. A. flavoviride 

Stipe and rachis black, scaly 36. A. acrocarpum 

Pinnae much wider 39. A. fraxinifolium 

Veins anastomosing. 

Frond bipinnate 37. A. esculentum 

Frond pinnate or simple. 
Base of pinnae cuneate. 

Axes brown, naked 38. A. fraxinifolium 

Axes black, scaly 39. A. Cumingii 

Frond usually simple, base cordate 40. A, cordifolium 

U A. MINUTUM Copel. Plate 39. 

A. minutum Copel., Univ. Calif. Publ. Bot. 18 (1942) 221. 

A. nanum, rhizomate brevi-repente, paleis linearibus 5 mm 
longis integris olivaceo-ferrugineis vestito; stipitibus fascicu- 

'^'* * Copeland: Aspidiaeeae of New Guinea 463 

latis, gracilimis, ca. 5 cm longis, fuscis, parce et decidue squa- 
mulatis; lamina 3-5 cm longa, basi 2.5 cm lata, subacuta, 
tripinnatifida; rhachi flliforme, superne angustissime alata, 
parce decidue strigillosa; pinnis ca. 8-paribus, inferioribus 
stipitulatis, apice rotundatis, infimis miaximis 8 mm latis, herba- 
ceis, glabris; pinnulis usque ad 4-paribus, infimis pedicellatis, 
truncato-orbicularibus, prof unde incisis segmentis 2-4, segmentis 
majoribus et pinnulis minoribus apice pauci-dentatis; soris ad 
pinnulas vel segmenta semper basalibus, saepe solitariis, bre- 
vibus, indusio lato, stramineo, integro vel eroso, persistente. 

Dutch New Guinea: Northern slope of Mt. Wilhelmina, alt. 
3,950 m. Brass et Myer-Drees 10107, in a rock-hole on grass- 
land; No. 108 iOy alt. 3,800 m, among mosses in wet limestone 

A relative of A. setiferum, distinguished by non-acuminate 
frond, ovate pinnae, dissected pinnules, and position of sori. 

2, A. SETIFERUM C. Chr. 

A. setiferum C. Chr., Index (1905) 146; Brittonia 2 (1937) 293. 

Brass et Myer-Drees 9857, Mt. Wilhelmina, alt. 3,560 m, in 
tufts in ground moss in subalpine forest; Brass 4892, H81, 
Mt. Albert Edward, alt. 3,680 m,— see Brittonia, 1. c. 

Himalayas. Represented in Borneo by A, atropurpureum 
Copel.; in Luzon and Formosa, by A. Oreopteris Copd. 

3. A. MYER-DREESU Copel. 

A, Myer-Dreesii Copel., Univ. Calif. Publ. Bot. 18 (1942) 222. 

A. rhizomate erecto, paleis f errugineis f uscescentibus integris 
anguste lanceolatis attenuatis 7 mm longis vestito; stipitibus 
fasciculatis, 30 cm altis, gracilibus, stramineis brunnescentibus, 
deorsum sparse paleatis sursum nudis; lamina 20 cm longa 10 
cm lata, subtripinnata, apice valde attenuata, glabra, papyracea 
(sicca) ; pinnis infimis 8.5 cm longis 15 mm latis, sensim acu- 
minatis, pedicellis 3 mm longis; pinnulis majoribus brevi- 
pedicellatis, rhomboideis, 10 mm longis 5 mm latis, apice rotun- 
datis, subpinnatis segmentis 0.6-1.2 mm latis apice 1-3-dentatis ; 
pinnis sequentibus gradatim minoribus; soris ad segmenta pie- 
risque basalibus ; indusiis latis, plerisque rectis, brunneis, integris 
vel eroso-crenatis. 

Dutch New Guinea: 7 km N.E. of Wilhelmina-top, alt. 2,560 
m, Brass et Myer-Drees 9851, in ground moss of subalpine 
forest. Type in Gray Herbarium. 

One of the group of A. nigripes, more immediately related 
to A. setiferum. 

108S2 7 

464 The Philippine Journal of Science ^^^^ 

4. A. SCOTINUM (Eos.) Copel.» comb. nov. 

Diplazium scotinum Ros. Fedde's Repert. 12 (1913) 169. 

The type is Kyesser B (or //) U, Mt. Bolan, alt. 3,400-3,800 
m; isotjrpe in Herb. Univ. Calif. Rosenstock, 1. c, distinguished 
also vars. platyloba, Keysser B 61 (not seen), and microloba, 
Keysser B 41, collected at the same place; the latter needs no 
distinguishing name. There is also var. contracta Hieron., 
Engler's Jahrb. 56 (1920) 143. 


5. A. MOROBENSE Copel., sp. nov. Plate 40. 

A. rhizomate adscendente, 3 mm crasso, paleis atris lanceo- 
latis 3 mm longis vestito ; stipitibus congestis, basin versus nigris 
paleatis, sursum rhachibusque atroviridibus glabris; lamina 
20-25 cm longa, 12 cm lata, bipinnata, acuminata; pinnis erecto- 
patentibus, pedicellatis, basi conspicue obliquis, acutis; pinnulis 
infimis acroscopicis maximis 15 mm longis, 7 mm latis, obliquis, 
latere acroscopico incisis lobis paucis, alibi cum pinnulis caeteris 
tantum serrulatis, papyraceis, glabris; soris majoribus 2 mm 
longis, plerisque unilateralibus, indusiis angustis obscuris. 

New Guinea: Morobe: Boana, alt. 800-1,400 m, Clemens 
U537, anno 1940. 

This species suggests the group of A. Williamsii, which is 
better developed in the Philippines. 

6. A. PEDICELLATUM Copel. Plate 41. 

A. pedicelMum Copel., Univ. Calif. Publ. Bot. 18 (1942) 222. 

A. rhizomate adscendente, 5 mm crasso, atro-fusco, apice 
paleis 1-2 mm longis ferrugineis vestito, glabrescente ; stipite 
10-15 cm alto, 1.5 mm crasso, nigrescente, glabro; lamina 25 
cm longa vel majore, ovata, tripinnata (plantarum minorum 
laminis jam soriferis tantum tripinnatifidis) , ad insertionem 
quamque pinnae vel pinnulae palea plerumque solitaria omata, 
alitor glabra, rhachibus atroviridibus, pinnulis et pinnulis fere 
recte distantibus pedicellatis ; pinnulis " oblongis, ca. 4 mm longis 
et 2 mm latis, integris vel apice denticulatis, herbaceis, venulis 
simplicibus ; soris usque ad 2 mm longis, fere omnibus unilatera- 
libus infimis rarius diplazioideis, indusio firmo, secus venulam 
nigro alibi laete brunneo. 

New Guinea: Morbe: Boana, alt. 800-1,400 m, Clemens 
U566; anno 1944. 

Related to A. latilobum, which has the same peculiar paleae 
at the nodes of the axes, but is distinguished by pubescent axes. 

'^'^' * Copeland: Aspidiaceae of New Guinea 465 

7. A. HUTTONI (Baker) Copel., comb. nov. 

Asplenium Hutt<mi Baker, Annals of Bot. 5 (1891) 311. 
Athyrium australe vax* papuanum C. Chr., Brittonia 2 (1937) 293. 
Diplazium cyatheifolium Ros. et al, in herb., non Presl. 

Brass 138^8, alt. 130 m; Brass 8863, Diene, Papua, alt. 500 m, 
type of var. papuanum; Carr H9i5, Boridi, Papua, alt. 1,200 m; 
Clemens (sent as host of a fungus), Morobe; from the same 
region, Schlechter 16800, Bamler 92a and 136, Bamler, Rosen- 
stock Fil. novog. exsicc. n. 221 . The species thus occurs at minor 
altitudes throughout New Guinea. It might hardly be identified 
as this species except by comparison with the type; I construe 
it by the Carr collection, received with this name from the 
British Museum (Mr. Alston). It was originally described 
from a specimen from the "Malay Isles," received by Baker 
from Messrs. Veitch. 

8. A. LATILOBUM Copel. Plate 42. 

A. latilobum Copel., Univ. Calif. Publ. Bot. 18 (1942) 222. 

A. silvatico (Blume) Milde affine, rhizomate breve adscendente, 
apice paleis nigrescentibus triangulari-ovatis acutis usque ad 
9 mm longis basi 3 mm latis vestito; stipitibus fasciculatis, 
30-35 cm altis 4 mm crassis, deorsum sparse appresso-paleatis 
alibi rhachibusque minute furfuraceis nigro-f uscis ; lamina 65 
cm longa, 40-45 cm lata, subtripinnata, pinnis infimis pauUo 
diminutis interdum deflexis, medialibus 20-25 cm longis, 8-10 
cm latis, acuminatis, basi pedicellatis (vix 1 cm) pauUo angus- 
tatis, sursum per pinnas pinnatas pinnatifidas serratas et 
integras ad apicem brevum acuminatum transeunte; pinnulis 
pinnarum medialium vix contiguis, 4-5 cm longis, ca. 16 mm 
latis, plerisque obtusis rarius acutis, basi truncatis pedicellatis 
(1.5-3 mm), deorsum pinnatis sursum profunde pinnatifidis; 
pinnulis" sessilibus et segmentis sequentibus decurrenti-con- 
fluentibus late oblongis, 4-5 mm latis, f alcato-truncatis, integris 
vel maximis infimis rarius late crenatis, atroviridibus, herbaceis 
vel siccis tenuiter papyraceis ; venis in segmentis pinnatis, venula 
infima acroscopica solummodo plerumque furcata; soris, infimo 
acroscopico saepe excepto, simplicibus, 2 mm longis, indusio 
f usco stramineo-marginato nigrescente. 

Dutch New Guinea: Idenburg River, alt. 850 m; Brass 13U20, 
type, "A large clump on shady bank of stream in rain forest; 
stipes and rachis black." Also, Nos, 12893, alt. 1,100 m, and 
12270, alt. 1,700 m. 

466 The Philippine Journal of Sdmwe ^^^^ 

The larger pinnules of the medial pinnae have usually a 
single pair of sessile secondary pinnules, sometimes a pair of 
adnata pinnules^ then about 4 pairs of similar but confluent 
segments. No. 12270 is a more lax specimen, the only one 
witii acute pinnules, and up to 7 pairs of segments and secondary 

This might be suspected of being Diplazium opadfolium 
V.A.V.R., Nova Guinea 14 (1924) 14, which I know by description 
only; but that specias is de^ribed as less compound, with 
narrower pinnules, and subcoriaceous ; and texture is usually 
a group characteristic among these ferns. 

9. A. 0ECOMP(^ItVM (Ros.) Coi»el. Plate 43. 

A, deeompositum (Ros.) Copel., Univ. Calif. Fubl. Bot. IS (1942) 222. 

Brachysorus, stipite ultra 60 cm alto, f usco, spinuloso, sparse 
furfuraceo; pinna inflma unica visa 32 cm longa, 20 cm lata, 
quadripinnatisecta, papyracea, rhachi f usca superne f urf uracea 
aliter glabra, pedicello 15 mm longo ; pinnulis imbricatis, media- 
libus 12 cm longis 3.5 cm latis, acuminatis, pedicellatis, recte 
patentibus vel recurvis, pinnulis inferioribus pauUo minoribus; 
pinnulis secundariis approximatis, infimis sessilibus, ceteris 
adnatis fere omnibus ala costale confluentibus, acutis, majoribus 
% costam versus incisis, segmentis multis 1 mm latis 
falcatis acutis; venis usque ad 4-paribus; soris costularibus 
(ad bases venarima), brevibus, indusio fusco parvo lacero. 

New Guinea: Sattelberg, alt. 800 m, Bamler 128 (1914), 
type in Herb. Univ. Calif.; received from Dr. Rosenstock as 
"Diplazium brevisorum J. Sm var. novoguineensis Ros. n. v., 
= Athyrium silvaticum Bl. f. decomposita Ros. n. f., an spec, 
nov.'^ This would give a correct idea of the affinity of the 
plant. However, the confusion is remarkable. "D. silvaticum 
Bl. var. novoguineensis Ros.'*, Hedwigia 56 (1915) 351, typified 
by Bamler 56 is segi*egate of D. silvaticum (Bory) Sw., correctly 
placed by Hieronymus, Engler's Jahrb. 56 (1920) 134; it is 
No. 32 A of this enumeration. 

10. A. WOODWARDIOmES (Presl) Christ 

A, woodwardioides (Presl) Christ, Verh. Nat. Ges. Basel 11 (1895) 

Brachysorus woodwardioides Presl. Epim. (1849) 70. 

Schlechter 1661S (alt. 300 m), 191H; Bamler, Rosenstock 
Fih novog. exsicc. n. 222, as Diplazium brevisorum J. Sm. I 
have no Cuming specimen for comparison and am not quite 
certain of the identification, made by Hieronymus for the 

'^'^'^ Copeland: Aspidiaceae of New Guinea 467 

Schlechter specimen. It is not in the Brause enumeration, ap- 
parently prepared, as to this genus, by Hieronymus. 

11. A. SYLVATICUM (Blume) MUde 

A. sylvaticum (Blume) Milde, Bot. Zeit. 24 (1866) 376. 

Bamler ISO, Sattelberg, alt. 800 m. This again bears label 
as var. novoguineensis Ros., n. v. It is quite identical with 
Java specimens. 

Java. Further range uncertain because our specimens are 

12. A. BLUMEI Copel. 

A. Blumei Copel., Philip. Jour. Sci. 3 C (1908) 294. 

Diplazium polypodioides Blume, Enum. (1828) 194. 

Athyrium fimbristegium Copel., Philip. Jour. Sci. 9 C (1914) 5. 

King 386 (type of A. fimbristegium), i59, Papua; Keysser H; 
Zahn 8. n., Sattelberg. Hieronymus omits this species from 
Brause's enumeration, but describes 5 new species, ''Subspecies 
D. polypodioidis" or "e turma D. polypodioidis." I have no 
authentic specimens of any of them, and can list them only 
for what they may be worth. Of these great ferns, such criteria 
as size of pinnule always vary from part to part of a single 
frond, and the forking of the veins varies with the size of 
the segment. The indusium of specimens I recognize as A. 
Blumei is fimbriate in Java and in New Guinea ; it is entire on 
specimens so named from Indo-China and Assam. 


Diplazium Nymani Hieron., Engler's Jahrb. 56 (1920) 186. 

The type is Nymxin 698, Sattelberg, alt. 900 m. Carr 15712, 
from Papua, comes with this name from the British Museum, 
and fits the description. 

14. DIPLAZIUM SeimADERI Hieron. 

Diplazium Schraderi Hieron., Engler's Jahrb. 56 (1920) 141. 
The type is Ledermann 12076, Schraderberg, alt. 2,070 m. 


Diplazium Sehleehteri Hieron., Engler's Jahrb. 56 (1920) 138. 

The cited specimens are Ledeirmann 11032, 11867, 11942; 
Nymxin 432; Schlechter U3S9; Hellwig 609, 613, 617, all from 
Kaiser-Wilhelmsland, alt. 900-2,070 m. 

468 The Philippine Journal of Science ^^^ 


Diplazium SehuUzei Hieron., Engler's Jahrb. 56 (1920 140. 

Schultze 260; Ledermann 12882; Warburg s. n. Brass 5165, 
Mafulu, Papua, alt. 1,250 m is identified as this species by 
Christensen; it does not fit the description, but may have been 
named by comparison with an authentic specimen. 


Diplazium Naumanni Hieron., Engler's Jahrb. 56 (1920) 137. 

Described from a collection by the Gazelle Expedition at Mac 
Cluer Bay. 

Diplazium prolongatum Eos., Meded. Kijks Herb. No. 31 (1917) 5. 

"A praecedente recedit pinnis secundariis haud acuminatis, 
segmentis s. pinnis tertiariis margine serratis, non rotundato- 
crenatis." The preceding species is D. pseudocyatheifolium, 
which has not acuminate pinnules, nor any secondary segments 
or pinnules, and is nowhere round-crenate. Comparison with 
something else may have been intended, but, as published, D. 
prolongatum is essentially a nomen nudum, 

Diplazium asperum var. subpolypodioides v.A.v.R. [D. sub- 
polypodioides v.A.v.R., Bull. Jardin Suit. No. XX (1915) 11] 
is listed by Hieronymus, p. 136, but I find no explicit authority 
for its occurrence elsewhere than in Sumatra. 

17. A. SQUAMULIGEBUM (Ros.: Hieron.) Copel. 

A. squamuligerum (Eos.: Hieron.) Copel., Jour. Arnold Arb. 10 

(1929) 178. 
Asplenium varians var. squamuligera Ros.j Fedde's Repert. 12 (1913) 

Asplenium squamuligerum Hieron., Hedwigia 61 (1919) 5; Engler's 

Jahrb. 56 (1920) 147. 
Athyrium Ramosii Copel., Philip. Jour. Sci. 38 (1929) 140. 
A. geophilum var. C. Chr., Suppl. Ill (1934) 41 (overlooking p. 38). 

The assignment of this plant to a genus has made more than 
enough trouble. The latest instance is by Christensen, Brittonia 
2 (1937) 293, "The referring of Asplenium squamuligerum 
Hieron. to Athyrium by Copeland is very problematical." This 
does not appear in Suppl. Ill p. 41, under A. geophilum, and my 
type sheet of A. Ramosii is annotated by Christensen as "cer- 
tainly an Athjrrium." Still, its relatives are plants which he still 
prefers to call Diplazium. With the fine new New Guinea 
material in hand, I must conclude that A. Ramosii is the same 
species. A. geophilum, however, is distinct, being without the 
characteristic paleae on the axes. 

^'^* * Copeland: Aspidiaceaeof New Guinea 469 

Keysser 228, Sattelberg Hinterland, alt. 1,400-1,500 m, type; 
Brass 12209, 12882, 13685, alt. 850-1,750 m, Idenburg River; 
Clemens, Morobe, alt. 900 m. 



Diplazium opacifolium v.A.v.R., Nova Guinea 14 (1924) 14. 

Lam lOiS, Mamberamo River, alt. 90 m; not seen. 

A collection by Zahn, Sattelberg, alt. 900 m, received with a 
manuscript name by Christ, fits the description of D. opacifolium 
in most respects, but the largest pinnules of the complete frond 
are less than 2 cm long by 8 mm wide. If it were a Philippine 
specimen, it might be A. atratum. 

19. A. NITENS (Ros.) Copel., comb. nov. 

Diplazium nitens Ros., Fedde's Repert. 5 (1908) 373, 

Werner 62, Danum, alt. 500 m; isotype in Herb. Univ. Calif.; 
Bamler, Rosenstock Fil. novog. exsicc. n. 161, Logaueng, alt. 
400 m; Schlechter 176^7, Kani Mountains, alt. 600 m. 


20. A. DAVAOENSE Copel. 

A. davaoense Copel., Philip. Jour. Sci. 3 C (1908) 295. 

Bamler, Rosenstock Fil novog. exsicc. n. 189, Sattelberg, 
alt. 800 m; Carr 18018, 1U71, Boridi, Papua, alt. 1,400 m; all 
received as Diplazium cyclolohum (Christ) Ros., a name which 
I have not located. 


21. A. MAXIMUM (Don) Copel. 

A. maximum (Don) Copel., Philip. Jour. Sci. 3 C (1908) 295. 

Both Diplazium maximum (Don) C. Chr. and D. latifolium 
Moore have been reported from New Guinea. I have no local 
specimen, and, as both names have been used for groups of 
species, do not know at all exactly what were (or is) meant. 

22. A. CYATHEIFOLIUM ORich.) Mildc 

A, cyatheifolium (Rich.) Milde, Bot. Zeit. 28 (1870) 853. 

King 381; Brass 603, 5S05; Papua. 
Endemic, so far as definitely known. 

23. A. UNCIDENS (Ros.) Copel. 

A. unddens (Ros.) Copel., Univ. Calif. Publ. Bot. 12 (1931) 394. 
Dryopteris unddens Ros., Fedde's Repert. 10 (1912) 337. 

470 Tf^^ Philippine Journal of Science ^^*® 

Brass 12894, alt. 1,100 m; Frau Bander 8, Sattelberg, isotype 
in Herb, Univ. Calif.; Keysser 17 9p, ibid., alt. 1,400-1,500 m; 
Clemens, Morobe District. Exindusiate. 


24. A. KEYSSERI Copel.^ nom. nov. 

Diplazium atropurpureum Ros., Fedde's Repert. 12 (1913) 528, non 
Athyriiim atropurpureum Copel. 

Keysser 179, Sattelberg Hinterland, alt. 1,450 m; isotype in 
Herb. Univ. Calif.; Brass 50U, Mt. Tafa, Papua, alt. 2,400 m. 

25. A. PROTENSUM (Ros.) Copel.» comb. nov. 

Diplazium protensum Ros., Fedde's Repert. 12 (1913) 169. 

Keysser 106, 108 (as published), Sattelberg, alt. 800-1,000 m; 
not certainly seen; but we have, from Dr. Rosenstock, Keysser 
II 118, slightly smaller but otherwise as described, probably 
different in number because of error. Most sori are exindusiate, 
but a few narrow indusia can be detected. Like the preceding 
species in its paleae and prior in name, but apparently distinct 
in other respects. 


Diplazium asterothrix C. Chr., Britonnia 2 (1937) 292. 

Brass 8848, Dieni, Central Division, Papua, alt. 500 m. 
Schkehter 16853, received as Dryopteris africana, is probably 
an ill developed form of the same species. 

This is like the two preceding species in its stellate paleae, 
and like A. uncidens in being exindusiate. I suspect that it 
is more perfectly developed specimen of A. protensum. 

27. A. SORSOGONENSE (Pxesl) Blilde 

A. sorsogonense (Fresl) Milde, Bot. Zeit 2B (1870) 354. 

Brass 8945, Cyclops Mountains, alt. 500 m; 18655, Idenburg 
River, alt. 700 m; Kmg 829; Carr 12071, 12545, Papua. 

I have not seen Diplazium mamberamense v.A.v.R., Suppls 
Corections (1917) 55, but suspect it to be an ill developed 
A. sorsogonense, distinguished by pinnae cut only half-way to 
the costae. 

To India; but slightly different beyond the Malay Peninsula. 

28. A. ARCHBOLDH Copel. Plate 44. 

A. Archboldii Copel., Univ. Calif. Publ. Bot. 18 (1942) 222. 

A. rhizomate adscendente, paleis brunneis anguste lanceolatis 
17 mm longis prime integris tum demum margine fibrilloso- 

'^'^'^ Copeland: Aspidiaceae of New Guinea 471 

deliquescente vestito; stipite 40 cm alto, sicco 4 mm crasso, 
superne trisulcato f usco, rhachique paleis nigrjs rigidis recurvis 
angustis 1-2 mm longis ornatis; lamina 75 cm longa, bipinna- 
tifida; pinnis iniimis 11 cm longis, 5 cm latis; medialibus 18 cm 
longis, 6 cm latis, acuminatis, basi truncatis vel abrupte paullo 
angustatis, breviter (1 nrni) pedicellatis, deorsum % ad costam 
pinnatifidis, apices versus tantum serratis, subcoriaceis, 
segmentis serrulatis acutis; pinnis superioribus etiam 15 cm 
longis ca, 3 cm latis vix i/^ ad costam incisis lobis rotun- 
datis; apice frondis pinnis coadunatis composito ca. 15 cm 
longo; venis plerisque furcatis; soris segmentorum ma jorum 
usque ad 10-paribus, ca. 6 mm longis, margine remotis, infimis 
diplazioideis, indusio f usco angusto integro. 

Dutch New Guinea: 15 km S.W. of Bernhard Camp, Idenburg 
River, alt. 1,700 m. Brass 12272; "Common ground fern in a 
rain forest; leaves somewhat fleshy." 

The upper part of the frond suggests A. maximum^ but the 
lower part is utterly different. It is probably a relative of 
Diplazium pseMdoshepherdioides, but the lower and medial pinnae 
af e twice as broad, and very short-stalked ; by description, that 
species seems much like A. polycarpum Copel., of Borneo. 

29. A. jrAPONICUM (Thunb.) Copel. 

A. japanicum (Thunb.) Copel., Philip. Jour. Sci. 3 C (1908) 290. 

Brass 9303, Lake Habbema, alt. 3,225 m, very large ; Clemens 
Morobe, ho3t of a fungus No. 6735, not quite identical. 
New Zealand (native?) to Japan and India, 

30. A. WEINLANDII (Christ) Copel., comb. nov. 

DipUmum Weinlandii Christ, Bull. Boiss. II I (1901) 452. 
P, bamlerianum Ros., Fedde's Repert. 10 (1912) 329. 

Bamler {No. ^0), Rosenstock Fil. novog. exsicc. n. 220, Sattel- 
berg, alt. 600 m, isotype in Herb. Univ. Calif. 

ai. A. CRENATO-SEERATUM (Blume) Milde 

A. crenatum'Serratumf (Blume) Milde, Bot. Zeit. 28 (1870) 853. 

King SU, U6. 
Malaya; Philippines. 

32. A. BUnuppraUM (Brack.) Copel. 

A. bumferum (Brack.) Copel., Bishop Museum Bull. No. 59 (1929) 

"4JJ2 The Philippine Journal of Science ^®*® 

Brass 8826, Hollandia, alt. 50 m. Common and variable. 
Most New Guinea specimens are notably large, 
Fiji to Mauritius. 


DiptaMum novoguineense (Ros.) Hieron., Engler's Jahrb. 56 (1920) 

Bamler 56 bis, Sattelberg, alt. 600 m, isotype in Herb. Univ. 
Calif., reissued as Rosenstock Fil. novog. exsicc. n. 219. This 
is the most over-grown development of A, bulbiferum, the pinnae 
up to 25 cm long and 4.5 cm wide, and very shallowly cut. It 
looks distinct enough for specific recognition, but probably 
intergrades with plants of more normal size. 


33. A. LEDERMANNI Hieroii. 

A. Ledermanni Hieron., Engler's Jahrb. 56 (1920) 133. 
Ledermann 11906, alt. 2,070 m. Not seen. 

34. A. PAIXmUM (Blmne) MUde 

A. pallidum (Blume) Milde, Bot. Zeit. (1870) 354. 
King S65, Papua; Bamler, Rosenstock Fil. novog. exsicc. n. 
187, Wareo, alt. 600 m. 
Malaya; Philippines; Queensland; New Hebrides. 

35. A. TLAVOVmiDE Alston 

A. flavovtride Alston, Jour. Bot. 78 (1940) 226. 

Clemens 70S7, Sambanga, Morobe, alt. 5,000-6,000 feet. Not 

36. A. ACROCARPUM O^os.) Copel., comb. nov. 

Diplazium acrocarpum Ros., Fedde's Repert. 10 (1912) 328. 
Asplenium acrocarpum Hieron., Hedwigia 61 (1919) 32; Engler's 
Jahrb. 56 (1920) 148. 

Brass 120S6, alt. 1,750 m, common ground fern in rain 
forest. Described from the Sattelberg, Keysser 27, isotype in 
Herb. Univ. Calif.; reported also as Ledermann 1185S. 


Report of A. Merrillii in New Guinea may be due to confusion 
with this species. The Philippine A. longissimum Copel. has 
far more numerous, smaller pinnae. The Bornean A. fvligi- 
nostvm has a long pinnatifid apex. These are all nearly related, 
and with several relatives pinnatifid throughout, might constitute 
a small segregate genus. 

3t. A. l^CULENTUM (Retz.) Copel. 

A. escuUntum (Retz.) Copel., Philip. Jour. Sci. 3 C (1908) 295. 

■^"^•^ Copeland: Aspidiaceae of New Guinea 473 

Brass 136^6, alt. 850 m. Common at minor altitudes through- 
out New Guinea, always with only the lower veinlets anastomos- 
ing. This form has been named Asplenium vitiense Baker and 
A. dietrichianum Luerssen, thus occupying the eastern end of 
the range of the species in which it is here included. The 
typical form, with half or more of the veinlets anastomosing, is 
found in Hawaii, but is probably not native there. 

To India. 

3S. A. FRAXINIFOLIUM (Presl) Milde 

A, fraxinifolium (Presl) Milde, Bot. Zeit. 28 (1870) 353. 

Brass 18881, alt. 120 m; Bamler 30, Logaueng; King 377, 
4>H, Papua. Schlechter 17793 bears this name, but is distinct. 
Also reported from New Guinea as Diplazium bantamense. 
Report of D. altemifolium may be based on this or the following 

To India and Formosa. 

39. A. CUMINGU (Presl) Milde 

A. Cumingii (Presl) Milde, Bot. Zeit. 28 (1870) 353. 

Brass 1227 U, alt. 1,750 m; Bamler 121, Wareo, distributed 
as A. fraxinifolium; Brass BiM, Papua, alt., 1,450 m. New 
Guinea specimens are less scaly than the Philippine type, but 
clearly nearer to it than to A. fraxinifolium. 

Philippines; Celebes. 

40. A. CORDIFOLIUM (Blame) Copel. 

A. cordifoKum (Blume) Copel., Philip. Jour. Sci. 3 C (1908) 300. 

Brass 12250, 13U8, 137 iO, alt. 570-1,750 m. Conmion. 
To the Malay Peninsula. 

29. DIPLAZIOPSIS Christensen 
D. JAVANICA (Blume) C. Chr. 

D. javanica (Blume) C. Chr., Index (1905) 227. 

Brass 12939, alt. 1,200 m. 
Samoa to Formosa and India. 


Key to the species 

Frond simply pinnate 1. C. prolifera 

Frond bipinnate 2. (7. spinulosa 

1. C. PROLIFERA (Lam.) Bory 

C. prolifera (Lam.) Bory, Voy. I (1804) 283. 

Athyrium accedens (Blume) Milde, Bot. Zeit. 28 (1870) 353. 

474 The Philippine Journal of Sdmee i^^^ 

Brass 13799, alt. 160 m; probably common, certainly conspic- 
uous, at minor altitudes throughout New Guinea. 
Fiji to Africa. 

2. C. SPfinJLOSA (Bliime) J. Smith 

C. spinulosa (Blume) J. Smith, Jour. Bot. 3 (1841) 409. 
Diplazium spinulosum Blume, Enum. (1828) 193. 
Diplazium paradoxum Fee, Genera (1850-1852) 214. 
Asplenium smithianum Baker, Syn. Fil. (1867) 245; Christ, Ann. 
Jar. Buit. 15 (1904) 122. 

Bamler 66, Sattelberg, alt. 800 m; Carr U526, 15688, Papua. 
Rosenstock, Hedwigia 56 (1915) 351, distinguished two varieties 
or forms by name, and entered a third name on one of our 
labels; all may apply to parts of a single frond. 

The apparent distribution of this species, unknown between 
Celebes and Ceylon, is remarkable. 


[Photographs of types, except Plates 6 and 29, which are photo^aphs of fragments 
of types.] 

Plate 1, Polystichum Archboldii Copel. 

2. P. Brasii Copel. 

3. P. muticum Copel. 

4. P. Myer-Dreesii Copel. 

5. P. cheilanthoides Copel. 

6a. Lomagramma angustipinna Copel. 
6b. Cyclosorus adenostegius Copel. 

7. Elaphoglossum habbemense Copel. 

8. E. Archboldii Copel. 

9. E. laticuneatum Copel. 

10. E. brunneum Copel. 

11. E. repens Copel. 

12. E. fuscum Copel. 

13. Ctenitis speciossima Copel. 

14. C. pulchra Copel. 

15. C. habbemensis Copel. 

16. Tectaria pubescens Copel. 

17. Lastrea platyptera Copel. 

18. L. petrophila Copel. 

19. L. Regis Copel. 

20. L. subdimorpha Copel. 

21. L. belensis Copel. 

22. L. ophiura Copel. 

23. L. crassa Copel. 

24. L. wantotensis Copel. 

25. L. armata Copel. 

26. Cyclosorus distinctus Copel. 

27. C. vestigiatus Copel. 

28. C. paripinnatus Copel. 

29. C. deltipterus Copel. 

30. C. subappendiculatus Copel. 

31. C. gregarius Copel. 

32. C. multiauriculatus Copel. 

33. C. protectus Copel. 

34. C. terrestris Copel. 

35. C. albosetosus Copel. 

36. C. strigosissimus Copel. 

37. C. riparius Copel. 

38. C. morobensis Copel. 

39. Athyrium minutum Copel. 

40. A. morobense Copel. 

41. A. pedicellatum Copel. 

42. A. latilobum Copel. 

43. A. decompositum Copel. 

44. A. Archboldii Copel. 



I Full. IP. .lorit. Sn., Vol. "k. No. i. 


tm'Ki.Axit: Asmn.wr.AK of New UrixKA.f | Pmi.ii*. .lorit. Sri., Vol. 78, No. 4. 


ropKi.AXi»: Ahi'Iiiiackak of Nku (, 

riXKA.I ll'tni.iF. Jont. Sri., Vol. 7s, No. l. 


roi'Kl.AXlK AKPIIllAt'KAK tH> Nkw ilVlWA 

IPiitijp. .loru. Sri., Vol. 7H, No. 1. 


(%>fi>;L.\\u: A8piiJiA<'K.\F OF New CJnxKA.I 

! Pun IP. ,hn II, 8fi.. Vol. 7S, No, 1. 



I PtHLip, Joru. Sri,. Vol., 7.s. No. L 


<N»rKi,AM): Ahpii«i.\«i-:,-\f, of Nku <iI'ixea,| 

I I'mijp. .lorn. Hri., Voi . 7s, M«i. 1. 



PLATE 6 A & B. 

C'oi'ELAXH: Asi'iitiArFAK (IF Nkw (IrfXFA.I 

|1*IIIL1F. J<H'|{. S(^l., Vt)I., 7,S, No. I, 


iivrlnmnmrinhv Arnohl ArUm:hm\. 

X«- 9QS3 L..I.BmvH ;\tig. I!l,t 

"In a euahioii of h«patl«s m\ erpoaod SraftCfi 
■ «f tree. 


r<ii«Ki.Axi»; AspiiHArKAK or Ni-.w (Jri\KA.| 

fl'iiiup. .lorn. Sti., Vol. Is, HiK 1. 

,««* ^* J)>*t^t* Hk.^ fitJin 


t'oi'i.i \\it: Asi'iiii \t'r \i or \'i u (,i w 

I'litf i!>. J.. I n, Sn.. \'.ii . 7n. Nt 




ui'M\\!i: Asriia \fi> \! .M- S} \\ <,i!si.' 

I PlliiJi'. .h»i K. Sri.. Vu! . 7v, .\o. I. 



Plant* ©I Owtefc W*w €«!«*» 

PLATE 10. 

I'ot'i-! \ Mi: ;\si*ii!! \fi A! t»i' Ni w car^iA. 

j PHH !|'. .hti li. Sri.. \'«.i . y-^, Xi 

F>LATE 11. 

iH'l I \\l>- \^f'llt| \« I \! t<l Kl \\ <i 

ll'Hinr. .lorn, ^i'u, Vol. T^, No. 1. 

' mm. 




PLATE 12, 

Hii ! \ Mc \:-l'lUt U'l Xi .M Xi \\ ilfwi 


PLATE in. 

! 'ol'l ! \ \|i; \>l*!!>l 

I fi'. Jot u. Sri.. \"<>i . 7n. Nil, 4. 


<'«»ii<:i..-\xi): Ahi'!iha('i-:a!-: <»!•• Niav <jI.'iM':.\, 

Piiiur. .lorit. Hi'!.. Vol.. 'l -, N*. 


I'oFiCi,.'! .\r>: Asi'iiHAi'KAK OF New (I 

ll'inii!'. JiH u. Sri., Vol. 7s, Ko, I, 

^-?>" ..-^f • '' 



PLATE 16, 

C%ii'KLANit: Asfii>iaCEaK fir Nku (JnXKA.I 

I Ffiiiji'. Jon:, Kn., Vol. 7s. Ni». -L 

PLATE 17. 

Copeland: Aspidiaceab gf N"ew Guinea. I 

irmtjp. rU}vii. Sci., Vol. Th, No, -f. 

ToFKLAN-U: AsFIDIArKAK OF Nkw (J|!L\KA.| fPlllI.lP. .loriL Sl'L. VOL. 18, No. 4. 

PLATE 19. 


IPhii.11% .loriE. 8ci.. Vtn . 7S, Nm 4. 

PLATE 20. 

CoPlliAND: ASliUlACEAl-: nF Nku OriXKA.l 

I l»nii.iF. .font. Sci„ Vc3L. 78, No. 4. 

'i-.i «C;* ^U^ .4^A*<-M-^ f'/d^:-: 

PLATE 21. 

<'<>M.!i.,\M>: AsrimACEAh. of Mi. w <;riM.:.' 

I l* JoriJ. i^vh, Vm. 7s, M«>. -1. 

■..*..r!» *t^>v<*-'^ 

PLATE 22. 

<*HFi-.i.AKi»: Asp!i»i,.\n-;AE of Nku (JrixFA.I 

! i»llil.!l\ .lot'E. .^Cl-, V»L. 7b, Nli 

PLATE 23. 

('iH'Kl.AXIt: ASF11>IA<1CAF i»F Nf-.W <Jri\'KA.| IPlHllI'. loi'U. Stl., V«)l.. Th, N<i 

l:,4C^ ' 

PLATE 24. 

f'orei.A.xii; Asi'ltJiAfY.AF, *w Ni;i\ (Jiixka. | | Pifiuf. Ji.ri:. Sri,, Vol.. 7s, Nd. 4. 


>n'Vh.\\n: AsriiiiArKAF of Nkw <J!:i\'ea,| 

I PlHLIl". JorK. SCL, V<u„ 7s, No. 4. 

PLATE 26. 

C'oPB^LANi): AsPiBiACEAE OF New Guinba.] [Philip. Joue. Sci., ¥c)l. 78, No. 4. 

PLATE 27, 

t"!oi'KLAXlK AsFlHiAIKAK OF HkW fjllNKA.! 

1 1*1111. tr. Joi'K. SVL, Vol. 1^, No, L 

PLATE 28. 

Ourn.AXii: Asi'iitiArKAi-, of Nkw (Jtimca.! |l»iiii.ii'. .I*n*u. 8rt.. Vol. Is, No, l 

PLATE 29. 

C'lU'I'Il.AN'!*; Asi»IIllA<'HAI-; OF Nl-;U (^fiXKA.I 

H'liiLir. .liM'ii, Sei., Vol. 7s, M<», l. 

PLATE 30. 

tj>FKiA\l>; AHPilHAi'KAl-, ^H•' Nf.W firLNKV.f 

|PllIl.ll\ .loriJ. St'L, Vol,. Tk^ 5iii 

PLATE 31. 

i'tH'Kt.wn: .\spiiti.i<"i'.\f-, <»r Nk\\ i.'rixi.A.l Jul i;. Si'i., Vol. 7>. Nv 

U * ' 

PLATE 32. 

foPEI.AMl: Asi'flilAfFAK i>F Nhw UriXFA.l 

I Philip. Joue. Sci., Vol. 78, No, 4. 

PLATE 33. 


t PifiiJi'. ,I«»fK, i^eu. Vol.. 7s, Mo. 4, 


■ '*^ 




CuvKiASUt AHFii»tAi!i';Ai'; <m<' Nku <;ri\E>' 

I Pill! H'. .}m:i\, Sn.. Voi. 7.H, No. I. 

PLATE 35. 

r.ol'KIAND: Ai^PlItlAOKAl-; itV }4V\\ C,Vl\h.\.l 

IFiHiii'. Jitrij. Sci.. Vol.. lb. Nil. I. 

.*.- =*4j 


rofl.LANln ASI'II.IAC'KAF Of Nhu lillXK^.I U'lULIP. JiU'li. SCi.. V«kL. iH, Mil, 4. 

PLATE 37. 

i'oi'i I \\-t.- Asrn=s\»'i \i -n :\\-\\ {, 

I'nn n\ Joi n. <.'5., Voi. Is, N> 

PLATE 38. 


r Philip, .loirfL Srr.. Voi,. 7«. N<i. 4. 

■'.t--rR-<4--vt.^..-%i >jti->-tt.t.«<;-f/n' ' <.*/vi*f"' 



fl;ir'.ar.f I luu^'v;?: 

PLATE 39. 

I o|M i \\!*; \^^i•Uli \|-1 A! ■ < |- \i X\ (I 

[Philip. Jour. Sci., ¥01:,. 78» N'o. 4. 

^4S^X i 

-'/ii..-n.-.. .,..«\*f.%-*.^ . v<-'-' 

PLATE 40. 

t^H'KI.-WU: .•\r^ril>IA(>;AK or Nl'W fJl'lMlCA. I 

IPim.ip, Joi'K. Sn., V<iL, TS, No. 4. 


PLATE 41. 

t'oPKLAX'ii: Aki'Idiackae <»r New (inixRA. . 

lI'iiiLip. Jon. Sci., ¥0L. 78, No. 4. 

PLATE 42. 

(."orEi.AMi: AsrUHM-b-.w iH- Ml'W V,i iSh\\,l 

Ifntf!!'. Jtiri:, Sn.. Vt.i.. 'IS, N»». -l. 

PLATE 4?.. 

Copei.and: ASPIDIACIAE OF Ne\¥ GurHEA.] 

[Philip. Jcii"k. Sn,, ¥oi . IR, M^n t. 

PLATE -14. 



Page 97, line 6 for Zenker, should read Zenker. 
Page 97, lines 6 and 11, for Benin, should read Bouin. 
Page 151, line 20, for E., should read F. 
Page 151, line 23, for EE., should read FF. 
Page 208, line 31, for A. gradle Fee, should read 19. A. Cle-^ 
Page 209, line 3 from bottom, H. Nepah., should read Fl Nepal 
Page 211, line 14, for dent., should read deut. 
Page 213, after line 19 and before 13a. A. Regw, insert the 
entire first paragraph from p. 214. 

Page 217, line 7 from bottom, for toyayanum, should read 
Page 217, bottom line, for Goadyi, should read Goadbyl 
Page 218, lines 9 and 10, for insiticum, should read insiticium. 
Page 219, line 27, should read 9,000 feet Clemens 12384^ and 
bis. The type is Keysser 11. 

Page 220, reverse lines 9 and 10 from bottom, and line 10, for 
Schroder's, should read Schrader's. 
Page 223, line 9, for Novoy, should read Novog. 
Pages 223 to 227, in the running head, for Asplemaceae, should 
read Aspleniaceae. 

Page 224, line 15, should read Fronds bipinnatifid or bipinnate. 
Page 225, in the running head, for Aspeneacea, should read 

Page 228, line 6, from bottom, for Bamher, should read Bamler 
Pages 233 to 274 were missed in pagination. 
Page 291, line 10, for (Hooker) ^ should read (Hooker) «. 
Page 292, line 1, for 8 to /* to 19 r/ , should read 8 to 10 ^ . 
Page 292, line 5, for Bismarch, should read Bismarck. 
Page 292, line 25, for dactic, should read lactic. 
Page 293, line 7, for PI. 1, fig. 2, should read PI. 1, fig. 1. 
Page 294, line 1, for fig, 2, should read figs. 1 and 2. 
Page 294, line 3, for PI. 1, fig. 5, should read PI. 2, fig. 5. 
Page 296, line 37, for 4 Helt, should read 4 Heft. 



[New names and new combinations are printed in italice,} 

A, 186. 188. 192, 195. 198. 

Acanthace®, 295. 

Achirophichthys Kampenl (Weber and de 

Beaufort), 152. 
Acrophorus Presl. 390. 891. 

Blumei Ching. 391. 

loxoscaphoides (Baker) Alston, 891. 

stipellatus (Wallich) Moore, 891. 

stipellatus var. montana Ros., 391. 
Acrostichum Linmeus, 36. 

aurenm L., 36. 
Addd, 170. 191. 
Adelmeria, 4. 
Adiantum Linnaeus, 39. 

caudatum L., 39. 

Christii Ros.. 39. 40. 

fulvum Raoul, 40. 

hispidulum Swartz. 39. 

hollandire V.A.V.R.. 39, 40. 

Kingii Ctopel.. 89. 

neo-guineense Moore, 89, 40. 

philippense L., 39. 
Adin (o) tS. 182. 
Adton, 181. 

kaan6 ti, 181. 
A^es. 46. 

Aegiceras corniculatum (L.) Blanco, 108. 
Ag. 168. 169. 170. 171. 203. 
Aga, 170. 
Agan>an6. 183. 
Agarfip. 198. 
Agdamfi, 178. 
Agin, 170. 
Aggap6. 201. 
Agiiitd, 203. 
Agingfina, 203. 
Agmalmalem, 174. 
Agnan&yon, 176. 
Agpa. 174, 203, 204. 
Agpaand. 183. 
Agpdda. 186. 
Agpap&n, 203. 

pay, 193. 
Agpatndg, 174. 
Agraman. 204. 
Agtufigpal, 203. 
Akin. 179. 

Alama ( Pangasinan ) , 95. 
Aliraango (Tagalog and Visayan), 95, 337. 

sa bntas, 96. 
Allek, 187. 

Alsophila dxyopteroidea Brause, 429. 
10882 8 

Amamaiilran, 187. 
Amangdn. 187. 
Amm6 kadi. 181. 

pay. 181. 
Ammok kadi, 181. 

pay. 181. 
Ampelopteris Kunze, 460. 

prelifera (Retz.) CJopel, 460. 
An, 169, 170. 
An-ano6n, 183. 

(kadi), 201. 
Ana ti, 182. 
Anid ritariid, 200. 

iflray, 200. 

pay, 200. 

ti, 182. 

ti gap6na, 182. 

ti ilim, 183. 
Anopheles, 44, 52, 55, 57, 58, 60. 64, 65, 69. 

filipinsB Manalang, 129, 181. 

mangyanus. 127-131. 

minimus flavirostria (Ludlow), 127-130. 

minimus var. flavirostris. 127. 129. 

(myzomyia) mangyanus (Banks). 129. 

(myzomyia) minimus flavirostris (Lud- 
low), 129. 
Anton kaand ti, 181. 
Antonand ti, 181. 
Antonii, Ilex. 2. 
Antono, 172. 
Apag, 169. 
Apagbalikas, 188. 
Apaganamd, 187. 
Apagap&man, 187. 
Apagdarikmdt, 188. 
Apagdipas, 188. 
Apagkanito. 188. 
Apagkigmdt, 188. 
Apdman, 187. 
Apay, 182. 
Apaya, 182. 
Apay-dpay, 182. 
Ari, 170. 
Arin, 170. 
Aristolochiaceie, 295. 
Arthrobotrya J. Smith, 401. 

articulata J. Smith, 401. 
Asclepiadeae, 295. 
Asin (o) ti, 182. 
Aspidium alatellum CJhrist. 445. 

bamlerianum Ros., 416. 

coadunatum Wall. var. rufovilkwa Ros., 




cliry»>tric}ium Baker, var. echinocarpa. 

chrysotrichum Baker, var. lobato-cre- 

nata, 412. 
devexum Kunze, 414, 
echinatum Mett., 433. 
esitensum Blume, 460. 
irrandifoliiim Presl, 417. 
Hispidulwn Decaisne, 455. 
Labmsca Christ, 415. 
membranaeeum Hooker, 414, 
nudum (Baker) Diels. 418. 
obliquatum Mett., 431. 
pfushyphyllum Kunze, 417. 
persoriferum Copel., 417. 
phaeostlgma C^ati, 412. 
profereoides Christ, 412. 
pteroides Sw., 467. 
quinquefoliolatum C. Chr. 418, 419. 
repandum WiUd., 417. 
subaequale Bos., 412. 
ulifirinosum Kunze, 437. 
vastum Blume, 414, 415. 
Warburgii Kuhn and Christ, 441. 
Aspleniaceae. 207. 
Asplenium Linnaeus, 207. 

acrobriruin Christ, 207, 211. 

acroearpum Hieron., 472. 

acutiuseulum Blume, 208, 217. 

afilne Swarts, 209, 220. 

amboitiense Willd., 208, 212. 

apoense, 212. 

Bakeri C. Chr., 17. 209, 222. 

bipinnatiildum Baker, 208, 214. 

Brassii C. Chr., 209, 220. 

resatianum Baker, 213, 215. 

Clemennm CopeL, sp. nov., 209, 216. 

Comosum Christ, 207, 211. 

cromwellianum Ros., 208, 217, 218. 

euneatum Lam., 209, 220, 221. 

cuneatum var, oceanicum Kuhn, 221. 

cymbifolium Christ, 207, 210, 212. 

decorum Kunze, 208, 214. 

deeorum Kimse var. acuminata Ros., 214. 

distans, 215. 

dubium Brack., 214, 215. 222. 

durum Copel., 207, 212. 

ellipticum (Fee) Copel., 209. 

ensiforme, 212. 

filipes Copel., 208, 213. 

Foersteri Ros., 209, 218. 219. 

GJellerupil v.A-v.R., 218. 

Goadyi Copel, and Shaw. 217, 218. 

irracile F£e, 216. 

hapalophylhmi Ros., 208, 219. 

Huttoni Baker, 465. 

impressivenium v.A.y.R.. 221. 

ineiso-dentatum Ros., 220. 

insimium Brack, 208, 216, 218. 

Kelelense Brause, 209, 221. 

Kefysserianum Bos., 208. 216. 

Kingii Copel., 213. 

laneedlatum Forsk., 209, 219. 

laserpitiifollum Lam., 209, 221. 

laxifolium V.A.V.R., 209. 220. 
Ledermanni Hieron., 217. 
lobulatum Mett., 217, 218. 
ludens Baker, 214. 

ludens var. dareoides Alston, 221, 222. 
macrophyllum Swartz, 208, 217. 
monotis Christ, 208, 214. 
moroben8e Copel., sp. nov., 207, 210, 211. 
multilineatum Hooker, 214, 215. 
multilineatum var, dareoides Ros., 221. 
nidiforme v.A.v.R., 209. 
Nidus L.. 207, 209. 
normale Don, 203, 212. 
novo-iruineense Ros., 209, 222. 
nutans Ros., 209, 221. 
paedisrens Copel., 209, 218, 220. 
papuanum Copel., 208, 212. 
paucidens v.A.v.R., 211. 
pellucidum Lam., 208, 216. 
persicifolium J. Smith, 208, 213. 215. 
Phyllitidus Don.. 207. 209. 2l0. 
planicaule Wall., 208, 216. 
praemorsum Swartz, 219. 
pseudoYuIcanicum v.A.v.R., 213. 214. 
Reffia Copel., sp. nov., 208, 213. 
remotum Moore, 208, 213. 214, 215. 
remotum Moore var. dareoides Ros., 214, 

remotlun var. dareoides 221, 222. 
Sancti-Christofori Christ, 208, 217. 
scandens J. Smith, 209, 221, 222. 
Schultzei Brause, 209, 222. 
scolopendrioides J. Smith, 210. 
setisectvmi Blume, 209, 220. 
Shawii Copel., 209, 221. 
simonsianum, 210. 
simplicifrons, 212. 
smithianum Baker, 474. 
squamulatum Blume, * 210, 218. 
squamuligerum Hieron., 468. 
subemarsrinatum Ros.. 208, 218. 
subs emarsrinatum, 214. 
tafanum C. Chr.. 209. 220. 
tenerimi Forster, 208, 214. 
tenerum var. Belangeri. 214, 
toyayanum Wall, 217. 
Trichomanes L., 208. 212. 
unilateral Lam., 208, 218. 
varians var. squamuligera Ros., 468. 
vittaeforme Cav., 207, 210, 211. 
Wemeri Ros., 207, 211. 
Athyrium Roth. 461. 

accedens (Blume) Milde. 478. 
acrocarpum Ros.. 462. 472. 
Archboldii Copel., 462. 470. 
atratum. 469. 

australe var. papuanum C. Chr., 465. 
atropurpureum Copel., 461, 468. 
Blumei Copel., 461. 467. 
bulbiferum (Brack.) Copel., 462, 471. 
cordifolium (Bl^ne) Copel., 462, 473. 
crenatum-serratum (Blume) Milde, 462, 



Cumin^i (Presl) Mllde. 462, 473. 
eyatbeifolium (Rich.) Milde. 462, 469. 
davaoense Cop«I., 462, 469. 
decompositum (Res.) Cope!., 461, 466. 
dietrichianum Luerssen, 473. 
esculentmn (Rete.) Copel., 462, 472. 
fimbristegrium Gopel., 467. 
flavoviride Alston 462, 472. 
fraxinifolium (Presl) Milde, 462. 473. 
fuliginofiom, 472. 
geopbilum var. C. Chr., 468. 
horisontale Ros.. 431. 
Huttoni (Baker) Copel.. comb, nov., 

461. 465. 

Huttoni (Baker) Copel. var. papuanum, 

japonicum (Thunb.) Copel., 462, 471. 
Keysseri Copel., nom, nov.. 462, 470, 
latllobum copel., 464. 
Ledermanni Hieron.. 462, 472. 
lonffissimum Copel., 472. 
maximum (Don) Copel.. 462, 469, 471. 
Merrillii, 472. 

minntvm Copel.. 461. 462. 
marobente Copel.. sp. nov., 464. 
Myer-Dreesii Copel.. 461, 463. 
nigripes, 463. 

nitens (Ros.) Ck)pel., comb. nov. 462, 469. 
Oreopterls Ck>pel.. 463. 
pallidmn (Blmne) Milde, 462, 472. 
pedlcellatum CJopel., 461, 464. 
polycarpum Ck>pel., 471. 
protensxma Ros.. 448, 470. 
Ramosii Copel., 468. 
scotinum (Ros.) Copel., comb, nov., 464. 
scotinum (Roe.) Copel., var. contracta 

Hieron., 464. 
scotinum (Ros.) Copel. var. microloba, 

scotinum (Ros.) Copel. var. platyloba. 

setiferum C. Chr., 461, 463. 
sorsoKonense (Presl) Milde. 462. 470. 
squamuliserum (Ros.: Hieron.) Opel., 

462. 468. 

eylvaticum (Blume) Milde, 461, 467. 

sylvaticum Blume var. novoguineensis 
Ros., 467. 

uncidcnd (Ros.) Ck>pel.. 462. 469, 470. 

wwdwardioides (Presl) Christ, 461, 466. 

Weinlaiidll (Christ) Copel., «wnb. nov., 
462, 471« 

Williamsil. 464. 
Attdn. 181. 

kaan6 ti. 181. 
Attonan6 ti, 181. 
Att6no, 172. 

AVERY, ARTHUR, see Hamm and Averv. 

Aw&n. 170, 191. 

Ay. 184. 

Aya. 184. 

Azorella selago Hook. 294, 295. 


Babaen, 204. 
Basi. 197. 

Baari met laeiSlgr, 197. 

Notes on Philippine mosquitoes. 
XV. The chaetotaxy of the pupj© 
and larvae of Tripteroides. 43. 
Banhawin. 96, 101. 

(S. Oceanlca), 100, 101, 105. 107. 
Bassit. 177, 191. 
Bauhinese, 295. 
Bolbitia Schott, 398. 

arguta (F6e) Chingf, 398, 899. 

heteroclita (Presl) Chinjr, 898. 

parva (Copel.) Chinar, 898. 899. 

quoyana (Gaud.) Chin?. 398. 

Taylori (Bailey) Ching, 398, 399. 
Bignoniaceie. 295. 
Biisr, 186. 
Biit. 177. 
Bin-ifir, 186. 
Blechnace«e, 223. 
Blechnum Linnaeus, 223, 227. 

Archboldil C. Chr., 224. 226. 

bamlerianum Ros.. 225. 

Brassi Copel.. 224. 225. 

decorum Brause. 224. 226. 

dentatum Diels. 223. 225. 

deorso-lobatum Brause. 223. 224. 

Faseri (A. Cunn.) Luerssen, 224. 227. 

finlaysonianum Wall.. 224. 

fluviatile (R. Br.) Lowe, 227. 

Fraseri (A. Cunn.) Luerssen var. no- 
voguineensis Brause, 227. 

Fraseri (A. CJunn.) Luerssen var. phil- 
ippinensis, 227. 

Hieronymi Brause, 224, 226. 

Keysseri Ros., 223, 226. 

latiusculum (v.A.v.R.) C. Chr., 224, 226. 

Iiedermanni Brause, 223, 224. 

membranaceum (Col.) Mett., 227. 

nisrropaleaceum Alston, 22S, 224. 

nudius Copel., 224, 226. 

orientate L.. 223, 224. 

papuanum Brause, 223, 225. 

pendulum Brause, 224, 225. 

revoltttum (v.A.v.R.) C. <3hr., 224, 225. 

saxatile Brause, 223, 224. 
BOEKENOOGEN, H. A. The carotene eon- 
tent of the fruit of Momordlca co- 
chinchinensis Spreng, 299. 
Botany. Leaflets of Philippine, t. 
Brachysorus woodwardioides Presl, 466. 
Bulik, 95, 9$, 

on the mode of action of naganol 
(Bayer 205) as a trypanocidal agent, 

Csesalpinieae (Bauhinia), 295. 



CaUipteria Bory» 473. 

prolifera {laaa,) Bory 473. 
8pintiI<Ma (Blmne) J. Smith, 478, 474. 
Campiiun argutum Copel., 899. 
heteroclitum Copel., 398. 
parvtan (Ck>peL), 399. 
quovanum Ck>pel., 398. 
Taykn-i Copel., 899. 
Campylosramma pteridifonnis V.A.V.R., 419. 
Canarium luzonicum (Blume) A. Gray, 111, 

112. 120. 
Canaritim ovatum, 112. 
Cancer serratiis Forskal, 99. 
Caryophyllaceae, 295. 
Caryophyllaceous, 294. 
Catnltis torazame Tanaka, 151. 
CentropriBtis pluerospiius Giinther 153. 
hirundaceus Cuvier and Valenciennes, 
Anomalous secondary srrowth in some 
roots of Hydrocotyle asiatica Linnaeus, 
Cheilanthes Swartz, 36. 

belensis Weatherby, 36, 37, 88. 
farinacea, 88. 
srisrantea Ces., 437, 
javensis (Willd.) Moore, 36. 
papuana 0. Chr., 36, 38. 
setigera Blmne, 436. 
tennifolia (Burm.) Swartz, 86. 
Chelidoperca hirundacea Boulengrer, 152. 
Chelidoperca hirundinacea (Cuv. and Val.), 

Chenopodiimi ambrosioides L., 278. 
Choerodon anchorage, 151. 

balerenaiSf New species, 149, 151. 
marffaritifems, 151. 
melanostisrma, 151. 
obliaracanthus, 151. 
schoenleini, 151. 
zamboangas, 151. 
zosterophorus, 151. 
Colocasia, 46. 
Combretacese, 295. 

L. DEE. Human milk studies: I The 
thimnine content of mature normal 
milk and beriberi milk, 878. 
Coniogramme F^e, 35. 

fraxinea var, Gopelandl Chr, 85. 
macrophylla (Blume) Hieron, 35. 
macrophylla var. undulata Hieron., 86. 
Convolvulacete, 295. 

leaflets of Philippine Botany, 1; 
Pteridacese of New Guinea, 6; Asple- 
nif4%se and Blechnacess of New 
Guinea, 207; Aspidiac^e of New Gui- 
nea, 886. 
Craspedodictsnun Copeland, 25, 26. 
ffrande Copel., 26. 
Schkchteri (Brause) Copel., 26. 

Crassulaceae. 294. 

CRUZ, AUREOO O. Composition of Philip- 
pine singkamas oil from the seeds of 
Pachyrrhizus erosus (Linn.) Urb.. 
Ctenitis Christensen, 408. 

alpina (Ros.) Ck»pel., 408; 409. 

dissecta (Foster) Copel., 408. 

habbemensis Opel., 408, 411. 

hypolepioides (Ros.) Copel., 408, 409. 

pulchra Copel., 408, 410. 

sagenioides (Mett.) Ck>pel., 109, 408. 411. 

Speciosissima Copel., 408, 409. 

vilis (Kunze) Ching. 408, 409. 

vilis (Kunze) Ching, var. microloba, 

viscosa, 411. 
Culcita Presl, 7. 

villosa C. Chr., 7. 
Culex pupie, 52. 
Currania Copeland, 437. 

oyamensis (Baker) Copel., 437. 
Cyathea atrispora Domin, 429. 
Cyclopeltis J. Smith, 897. 

novoguineensis Ros., 897. 
presliana (J. Smith) Berkeley, 397. 
Cyclosoras Link, 438, 441. 

adenostegius Copel., 438, 442. 
alatellus (Christ) CJopel., 425, 438, 

448. 445, 447, 
Albociliatus Copel., 438, 443. 
albosetosus Copel., 440, 455. 
amboinensis (Willd.) Copel., 440, 444, 

appendfculatus (Blume) Copel., 448, 

aquatilis Copel., 440. 
Archboldii (C. Chr.) CJopel., 440, 452. 
arfakianus (Baker) Copel., 439, 450. 
aridus (Don) Ching, 440, 452. 
Atasripii (Ros.) Copel., 439, 451. 
austerus (Brause) Opel., comb, nov., 

438, 447. 
beccarianus (Ces.) Copel., 441, 460. 
biauritus, 448. 

canescens (Blimie) Opel., 441,460. 
canescens var. nephrodiiformis Christ, 

cataractorum Wagnerand Grether, 438, 

confertus (Brause) Opel., 440, 455, 

crassifolius, 445. 

cuspidatus (Blume) Opel., 440, 458. 
cyatheoides (Kaulfuss) Farwdl, 452. 
deltipterus Opel.. 439, 449, 450. 
dentatus (Forsk) Ching, 454. 
dichrotrichus Opel, 440, 455. 
dimorpha (Brause) Opel., comb, nov., 

438, 442, 443. 
dimorphus (No. 6). 459. 
distinctus Opel., 438, 444. 
extensus (Blume) Ching, 489, 450. 



farinosus (Brause) Copel., comb, nov., 

489. 446. 
glandulosus (Blume) Chingr, 440, 458. 
S^ingylodea (Schkuhr) Link, 440, 458. 
Sregarius Copel., 489, 451. 
hastato-pinnatus (Brause) Copel., 441. 

458, 458. 
heterocarpus (Blume) Ching, 489, 

446, 447. 
hispidulus (Dec.) Copel., 440, 455. 
imponens (Os.) Copel.. 440, 457. 
interruptus (Willd.) Ching, 440, 457. 
invisua (Forster) Copel., 440, 457. 
lanceolus (Christ) Copel., 441, 460. 
lonsissimus 444. 
malodonis, 448. 
megaphylloides (Eos.) Copel., 439, 

mesraphylloides (Ros.) (Jopel. var. gla- 

brescens Ros., 450, 451. 
megaphyllus (Mett) Chingf, 450. 452. 
megaphyllus (Mett.) Ching var. ab- 

breviata Ros., 452. 
micans (Brause) Copel., 448. 
morobensis Ck>pel., 441, 469. 
mutabilis (Brause) Ck>pel., 438, 443. 
multiauriculatus Copel., 440, 453. 
oblongus (Brause) Copel., comb, nov., 

489, 447. 
obtuslfoUus (Ros.) Copel, 488, 442. 
parasiticus (L.) Farwell, 488, 444, 
parapinnatus Copel., 439, 448, 449. 
pennifferus, 449. 

pentaphyllus (Ros.) Copel.. 441, 459. 
philippinensis, 457. 
protectus Ck>pel., 440, 453. 
rigidus (Ridley) Ck)pel., comb, nov., 

440, 462. 
riparia (No. 53). 448. 
riparius Opel., 429, 456. 
roemerianus (Ros.) Copel., 438, 441. 
sogerensis ((5epp) (3opel., 489, 450. 
sp., 447, 448, 451. 
strigosissimus CJopel., 440, 456. 
subappendlculatus Ck>pel., 489, 449. 
subpubescens (Blume) Ching, 464. 
supraspinigerua (Ros.) Copel., 439, 447. 
tadavensis, 450. 
terrestris CJopel., 440, 454. 
triphyllus (Sw.) Ck)pel., 441, 459. 
truncatus (Poiret) Farewell, 439, 452. 
unitus (L.) Ching, 489, 452. 
urophyllus (Wall.) Copel., 440, 458. 
urophyllus (Wall.) Copel., var. peraspera 

V.A.V.R., 455. 
vestigiatus C5opel., 489, 446. 
Warburgil (Kuhn and Christ) Wagner 

and Grether 488, 441, 443. 
WoUastonii (vJl.v.R.) Copel., 452. 
Cystodium J. Smith, 7. 

florbifolium (J. E. Smith) J. Smith, 7. 

Cystopteris Bernhardi, 461. 

tenuisecta (Blume) Mett, 461. 


Da, 195. 

Daddtima, 176. 

Dalpada Amyot et Serville, 286. 

DandanI, 177. 

Daphne iulia, 295. 

Dards, 189. 

Darea, 219. 

Darea Belangeri Bory, 214. 

Datdaytfi, 191. 

Datdayt6y. 191. 

Datura alba Nees, 281. 

fastuosa Linnseus, 281. 
Davallia amboynensis Hooker, 23. 

blumeana Hooker, 17. 

cicutarioides Baker, 9. 

Forbesii Gepp, 889. 

intramarginalis Cesati, 23. 

lanceolata Baker, 17. 

longipinnula Cesati, 23. 

moluccana Blume, 23. 

Novae-Guinese Ros., 14, 

stenoloba Baker, 23. 

and Dee. 
Dennstaedtia Bernhardi, 10. 

acuminata Ros., 9. 

ampla (Baker) Bedd, 18. 

articulata Ros., 12. 

concinna Ros., 10, 11, 18, 14. 

cuneata (J. Smith: Hooker) Moore, 
10, 11. 

flaccida (Forster) Bernhard. 10, 11. 

glabrata (Cesati) C. Chr., 10, 12. 

magnifica Copel., 10, 11. 

moluccana (Blume) Moore, 10, 12. 

Novse-Guineee, 10. 

novoguineensis, 10. 

novogwCneensia (Ros.) Copel., comb, 
nov. 12. 

papuana, 10. 

penicillifera v.A.v.R., 10. 

resinifera (Blume) Mett, 10, 12. 

Rosenstockii v.A.v.R., 12. 

samoensis (Brack.) Moore, 10. 

scandens (Blume) Moore, 10, 12. 

Shawii Copel. 10, 11. 

Smithii, 12. 

Smithii var. novoguineensis Ros., 12. 
Di, 180, 181. 
Diacaipe Blume, 890, 

aspidioides Blume, 890. 
Dicksonia L'Heritier, 5. 

Archboldii Copel., 5, 6. 

erythrorachis Christ, 12. 

glabrata (Cesati, 12. 

grandis, 6, 6, 

Hieronymi Brause, 5, 6. 

Ledermanni Brause. 5, 6. 

papuana F.V.M.. 7. 



rhombifolia Baker, 12. 

Sclil«eht«ri Bratise, 5, 6. 

sciurns C. Chr., 5, 6. 
Dictyopteria heterosora (Baker) Bedd., 419. 

pentaphylla V.A.V.R., 418. 
Didymoehlaena Pesvauic* 397. 

truneatula (Swartz) J. Smith, 397. 
Biplaziopsis Christensen, 473. 

javanica (Blume) C. Chr., 473. 
Diplazium acrocarpum Bos., 472. 

altemifolium, 478. 

aspenmi var, subpolypodioides v.A.v.R., 

asterothrix C. Chr., 462. 470. 

atropurpurem Ros., 470. 

bamlerianimi Bos., 471. 

bantamense, 473. 

brevisorum J, Sm., 466. 

cyatheifolimn Bos.. 465. 

cyclolobmn (Christ) Bos., 469. 

latifolium Moore, 469. 

mamberamense v.A.v.B., 470. 

Naumanni Hieron.. 468. 

nitens Bos.. 462, 469. 

no70gtUneense (Bos.) Hieron., 472. 

Nymani Hieron., 461, 467. 

opacifolium v.A.v.R., 462. 466, 469. 

paradoxum F^, 474. 

polypodioides Blume, 467. 

prolongattmi Bos.. 468. 

pseudocyatheifolium. 468. 

pseudoshepherdioides, 471. 

Schlechteri Hieron., 461, 467. 

Schraderi Hieron., 461, 467. 

SchtUtaei Hieron., 461. 468. 

scotinnm Bos., 461. 

silvaticum (Bory) Sw., 466. 

spinnlostim BImne, 474. 

Weinlandii Christ, 471. 
Bitdy. 179. 

Doryopteris papuana Copel., 38, 39. 
Bryopolystichum Copeland, 412. 

cyclosortis v.A.v.B., 412. 

Kingii Copel., 412. 

phaeostigma (CJesatl) Copel., 412. 

tamatana C. Chr., 412. 
Dryopteris J. Smith. 88. 

adenosteiria Copel., 442. 

africana. 470. 

alatella. 445. 

albociliata Copel., 443. 

albosetosa Copel., 455. 

alpina Bos.. 409. 

anffosta Copel., 455. 

aquatilis Copel.. 443. 

arbuseiila. 451. 

Archboldil C. Chr., 452. 

arfakiana (Baker) C. Chr., 443, 451. 

armata Bos.. 457. 

Atasripil Bos.. 541. 

atrispora C. Chr., 429. 

atrispora var. varieVestita C. Chr., 429. 

auatera Bra\ise. 446. 
bamleriana Bos.. 407. 
baaisora Copel., 434. 
bipinnata Copel.. 422. 
Brassii C. Chr., 423. 
brunneo-villosa C. Chr., 437. 
canescens var. incana Bos.. 460. 
canescens var. nephrodiiformis Christ, 

canescens var. novosuineensis Brause, 

cesatiana C. Chr., 460. 
chlamydophora Bos., 434. 
concolor (Langsd. and Fischer) Kuhn, 

conferta Brause, 455. 
conterminoides C. Chr., 420, 424. 
crassa Copel., 435. 
crassifolia var. purpureo-lilacina C. 

Chr.. 432. 
debilis (Mett.) C. Chr., 441. 
deltiptera Copel., 449. 
depauperata Copel., 441. 
diapbana Brause, 427. 
dichrotricha Copel., 455. 
dichrotrichoides v.A.v.B., 455. 
didymosora (Parish) C. Chr., 444. 
dimorpha Brause. 442. 
discophora Bos.. 897. 
distincta Copel., 444, 
Dryander, 404. 
echinata C. Chr., 433. 
ensrleriana Brause, 427. 
falcatipinnula Ck>pel., 433. 
farinosa Brause, 447. 
flavio-virens Bos., 423. 
finlsterrae Brause, 421, 426. 
fulerens Brause, 432. 
genuflexa Bos. 437. 
srlaucescens Brause, 421, 426. 
habbemensis Clopel., 412. 
hastato-pinnata Brai2se, 458. 
hecterocarpa, 447. 
hirto-pilosa Bos., 439, 446. 
faispida Brause, 442. 
hispiduliformis C. Chr., 438, 445, 450« 
horiasontalis v.A.v.B., 422, 482. 
hunsteiniana Brause, 436. 
hypolipioides Bos., 409. 
imponens (C^ati) C. Chr., 457. 
intermedia (Blume) O.K., 409. 
keysseriana Bos., 430. 
lanceola (Christ) Copel.. 460. 
Lauterbachii Brause, 431. 
Xiedermanni Brause, 431. 
Ledermanni Brause, 412, 
leueolepis, 437. 
logavensis Bos., 445. 
longissima (Brack.) var. novogui- 

neensis Bos.. 444. 
mamberamensis v.A.v.B., 460. 
marattioides Alston, 422. 
megaphylloides Bos.. 450. 451. 



micans Bratise, 458. 

mixta Ros., 426. 

mollis (Jacq.) Hieron.» 444, 454. 

morol)€n8i8 Cope!., 459. 

multiaurictilata Gopel.. 453. 

munda Ros., 426. 

muricata Brause, 448. 

mutabilis Brause, 457. 

nephrolepioides C. Chr., 436. 

notabilis Brause 429. 

novoiraineensis Brause, 426. 

oblanceolata Copel., 460. 

oblonsa Brause, 447. 

obtusifolia Ros., 442. 

oligolepia v.A.v.R., 421, 427. 

ophiura Copel., 434. 

paleacea (Swartz) C. Chr., 407. 

pallescens Brause, 426. 

papuana (Nephrodium dissitifolium Ba- 
ker). 407. 

paraphysata Copel., 448. 

paripinnata Copel., 448. 

Peekeli v.A,v.R., 422. 431. 

pentaphylla Bos., 549. 

perpilifera v.A.v.R., 488, 445, 455. 

perpubescens Alston, 424. 

Petelotii, 407. 

petrophila Copel., 424. 

phseostigma C. Chr., 412. 

pilose-squamata v.A.v.R., 440, 452. 

prolifera C. Chr., 460. 

protecta Copel., 453. 

pulchra Copel., 410, 

purpurascens, 408. 

pseudoparasitica v.A.v.R., 407. 

pseudostenobasis Copel., 425, 445. 

quadriaurita Christ, 421, 480. 

quadriquetra v.A.v.R., 421, 431. 

Reiris Copel., 425. 

repandula v.A.v.R., 442. 

riffidifolia v.A.v.R., 422, 431. 

riparia Copel., 457. 

roemeriana Ros., 441. 

rufo-pilosa Brause, 458. 

sasrenioides, 411, 412. 

sas^ttifolia, 451. 

schizophyila v.A.v.R., 428. 

Sehlechteri Brause, 428. 

sogerensis Gepp. 450. 

sepikenais Brause, 438, 443, 450. 

setigera O. K. 436, 437. 

sparsa (Ham.) O.K., 407. 

spe«iosissima Copel., 409. 

apinosa Copeh, 436. 

stellato-pilosa Brause, 421, 428. 

stenobasis C. Chr., 442. 

stereophylla V.A.V.R., 422, 431. 

strigosissima Copel., 456. 

subappendiculata Copel., 449. 

subarborea (Baker) C. Chr., 407, 408. 

subarborea (Baker) C. Chr., var. aten- 
iiata keysser, 408. 

subarborea (Baker) C. Chr. var, bifor- 
mia Keysser, 408. 

subarborea (Baker) C. Chr. var. 

decompofiita Keysser, 408. 
subarborea (Baker) C. Chr. var. quadrt- 

pinnata Keysser, 408. 
subattenuata Ros., 425. 
subdigitata Brause, 392. 
subdimorpha Opel.. 432. 
subxiigra Brause, 427. 
supraspinigera Ros., 447. 
suprastrigosa Ros., 446. 
tamieusis Brause, 439, 451. 
terrestris Copel., 454. 
uliginosa (Kunze) C. Chr., 437. 
uncidens Ros., 469. 
urophylla var. cuspidata, 458. 
vestigiata Copel.. 446. 
villosipes Gepp, 421, 427. 
wantotensia Copel., 436. 
wariensis Copel,, 428. 484. 


Ectenus Dallas, 285, 286. 

aereus Stal. 287, 288, 289. 

brevirostris Stal, 285, 286, 289. 

generosus Stal, 285, 287. 

mesoleucuc Bergroth, 285, 287. 

pudicus Stal, 285, 286, 288. 

spectabilis (Burmeister). 286, 287, 288. 

Spinostut sp. nov., 286, 287. 288. 
Elaphoglossum Schott, 402. 

angulatum (Blume) Moore. 403, 405. 

Archboldii Copel.. 403, 404. 

Blumei J. Smith, 404. 

bolanicrcum Ros., 402, 404. 

Brassii C. Chr.. 404. 

brunneum Copel., 403, 405. 

callifolium, 406. 

conforme, 405. 

Copelandii Christ, 404. 

Cumingii (Fee) Moore var. papuanum 
C. Chr., 403, 404. 

fuscum C^pel., 403, 405. 

habbemense, Copel., 402, 403. 

hellwigianum Ros., 402, 403. 

laticuneatum Copel., 403 405. 

novoguineense Ros., 403, 404. 

petiolatum (Swartz) Urban, 403, 404. 

repens Copel., 403, 406. 

sclerophyllum v.A.v.R., 403, 405. 

sordidum Christ, 403, 404. 
Elmer, Adolf Daniel Eduard, 1. 
Elmeri, Eugenia, 4. 

Ficus, 4. 

Quercus, 4. 
Elmeria, 4, 
Elmerina, 4. 
Elmerinula, 4. 
Elmeriobryum, 4. 
Ek)dea, 277. 
En, 169. 170. 175, 189. 
Enumeratio Hemlpterorum Stal, 289. 
Scylla (Crustacea: Portunidae) I. Re- 
vision of the genus, 95; II. compar- 



ative studies on spermatogenesis and 

oogenesis, 801. 
Eublechnnm, 225. 
Enparyphinm ilocanmn, 882. 
Euphorbia hirta Unnaeus. 281. 


SANTIAGO. A supplementary suide 
for rapid identification of Anopheles 
minimus flavirostris (Ludlow) and 
Anopheles mangyanus (Banks), 127. 

Gayam, 199. 

Giim, 200. 

Goniopteris rigida Ridley, 452. 
rudis Ridley, 458. 

Gymnapistes niger (Cuv. and Val.), 153. 

Gymnogramme palmata Baker, 419. 


Halaelurus rudis Tanaka, 151. 

torazame Tanaka, 161. 

Philippine solar salt industry, 85. 
Hetnigramma Christ, 419. 

grandifolia Copel.. 420. 

Hollrungii (Kuhn) Copel., 419. 
Hemiptera insularum philippinarum Stal 

Ilemipteris Rosvnstock, 35. 
Hemipteris Wemeri Ros., 85. 
' Hemuinitis prolifera Retz., 460, 
HERRE, ALBERT W. C. T. A new la- 
brid and other interesting Philippine 
fish records, 149. 
Heterogonium Presl, 412. 

profereoides (Christ) Copel., 412. 
Heteroneuron argutum F€e, 899. 

Naumanni Kuhn, 899. 
IHistiopteris (Agardh) J. Smith, 29, 31. 

estipulata V.A.V.R., 29. 30. 

incisa (Thumb.) J. Smith, 29, 30. 
Hydrocotyloideae, 294. 
Hydrocotyle asiatica L., 291, 294, 295. 
Hjrpolepid Bemhardi, 27. 

aculeata Gepp, 28. 

Archboldii Copel., 27, 28. 

bamleriana Ros, minor, 27. 

bamleriana f. minor, 27. 

distans, 28. 

grandifolia (]repp, 27, 28. 

grandifrons €repp, 10. 

papuana Bailey, 12, 27. 28. 

punctata (Thumb.) Mett., 27, 28. 

punctata (Thumb.) Mett. var. obscura 
Brause, 27. 

tenuifolia (Forster) Berhhardi, 27, 28. 

I, 169, 170, 202, 203. 


Ididy, 178, 180. 

Ikapi, 170. 

Ikapin, 170. 

In, 174. 

Ingg&, 203. 

Inggand, 203. 

Inn, 169. 

Intonand ti, 181. 

Intdn, kaand ti, 181. 

Intdno, 172. 

Ipa, 170, 204. 

Ipi, 170. 

Ipin. 170. 

Isoloma J. Smith, 24. 

fuligineum Copel., comb, nov., 24. 
Istdy, 187. 
Ita. 171. 
Itattd, 171. 
Itdy, 171. 
Itattdy, 171. 
Ithycaulon Copel., 7. 
1 acuminatum Copel., 9. 

cicutarioides Alston, 9. 

moluccanum Copel., 8. 

minus C. Chr., 8. 

Novae-Guinete (Ros.) Alston, 9. 

tenuisectum C. Chr., 9. 

Iti anifl ti, 181. 


Ka, 168, 169, 203. 

Ka. . . an, 168, 173, 203. 

Kaand ti, 181. 

Kaan6nto, 182. 

Kad, 198. 

Kadi, 198. 

Kalkala (pay), 188. 

Kandyon, 176. 

Kafigrondan, 187. 

Kanika, 192. 

Kapag, 169. 

Kara, 169. 

Kasand ti, 183. 

Kast&, 186. 

Kast6y, 186. 

Ket, 195. 

Ketdi, 195. 

Koma, 198. 


La, 186, 190. 

Ldeng, 186, 190. 

Lastrea Bory, 420, 423, 438. 

alta (Brause) Copel., comb, nov., 429. 

armata Copel.. 433. 

Beddomei (Baker) Bedd., 420, 423. 

belensis Copel., 422, 433. 

bipinnata Copel., comb, nov., 420, 422. 

Brassii Copel., comb, nov., 420, 424, 425. 

calcarata (Blume) Moore, 422, 432. 

chlamydophora (Ros.) CJopel., 422, 434. 

conterminoides Copel., comb nov., 420 

coriacea (Brause) Copel., comb, nov 
421, 428. 

coriacea (Brause) Copel., var. data 

costulisora Copel., 422, 434. 



craasa Copel., 422, 435. 

crassifolia, 488, 434. 

diaphana (Brause) Copel., comb, nov., 

421. 427. 
dryopteroidea (Brause) CTopel., 429. 
echinata (Mett.) Ck)pel., 422, 483. 
engleriana (Brause) CJopel., 421, 427. 
engleriana (Brause) dopel., var. hirta 

C. Chr., 427. 
ensipinna (Brause) Gopel., 421, 430. 
ensipinna (Brause) CJopel, var. acumi- 
nata, 430. 
erubescens (Wall.) CJopel., 428. 
falcatipinnula Copel., 422, 433. 
FinisterwB (Brause) Copel., 421, 426. 
flavo-virena (Brause) Copel., comb. 

nov., 422, 432. 
arracilescens (Blume) Hooker, 422, 431. 
Hallieri, 434. 

Harveyi Carr., 420, 425. 
Hunsteiniana (Brause) Copel., 422, 486. 
immersa (Brause) Moore, 421. 430. 
keysseriana (Ros.) Cope!.. 421, 480. 
Klossii Ridley, 428. 
Lauterbachii (Brause) Copel., 422, 431, 
leucolepis Presl, 487. 
marattioides (Alston) (3opel., 420, 422. 
mixta Copel., comb, nov., 421, 426. 
motleyana (Hooker) Copel., 433. 
nephrolepioides (C. Chr.) Opel., 422, 

426, 436. 
notabilis (Brause) Copel., 421, 429. 
novoguineensis (Brause) Copel., 422, 426. 
obliquata (Mett.) Copel., 422, 481. 
ophiura Copel., 422, 434. 
pallescens (Brause) CJopel., comb, nov., 

421, 426. 
perpubescens (Alston) Copel., comb. 

nov.. 420, 424. 
petrophia Copel.. 420, 424, 441. 
platyptera Copel.. 420, 423. 
pseudostenobasis Copel,, 420, 425. 
pyrrhorhachis, 428. 

quadriaurita (Christ) Copel., 421, 430. 
Regris Copel., 420. 425. 
setigera Bedd., 436. 
setosa Presl, 436, 437. 
spinosa Newman, 436. 
stellato-pilosa (Brause) CJopel., comb. 

nov., 428. 
subattenuata (Ros.) Copel., 420, 425. 
subdimorpha CJopel., 432. 
subnigra (Brause) Copel., 421, 427. 
tuberculata (Ces.) Copel., 421, 428. 
tuberculata (Ces.) Copel. var. djamuen- 

sis Brause, 428. 
varievestita (C. Chr.) Copel., 429. 
viscosa J. Smith, 421, 428, 426. 
wantotensis Copd., 422, 436. 
wariensis Copel., 421, 428. 
Lepidocaulon Copel., 80. 
caudatum Copel. 80. 

Leptochilus cuspidatus (Presl) var., mar- 
ginalis Ros., 899. 

novoguineensis Brause, 401. 
Leptogramma, 428. 
Leptolepia Mettenius, 26. 

Novse-Zelandiae (Col.) Kuhn, 26. 
Leucostegia loxoscaphoides, 892. 
Lindsaea Dryander, 15. 

adiantoidea (Blume) Kuhn, 15, 18, 19. 

azurea Christ, 16, 21. 

azurea Christ var. Mamb® V.A.V.R., 21. 

Bakeri C. Chr.. 15. 17. 

blanda Mett.. 16, 18. 

blumeana (Hooker) Kuhn, 17. 

brevipes Copel., 15, 19. 

concinna J. Smith, 15, 19. 

crassipes Ros.. 15. 19. 

cultrata, 19. 

davallioides Blume, 16, 22. 

decomposita Willd., 16, 20. 

ensifolia Swartz, 16, 22, 81. 

foersteri Ros., 15, 17. 

furcata CJopel., 16, 21. 

gracilis Blume, 15, 18. 

heterophylla Dry., 16, 22. 

hymenophylloides Blume, 15, 17. 

Kingii Copel., 15, 19. 

Ledermanni Brause, 17. 

macraeana, 16. 

marginata Brause, 15, 18, 21. 

marginata Brause var. falcata, 18. 

Merrillii, 16. 

microstegia Copel., 16, 20. 

monocarpa Ros., 19. 

monosora Ros., 19. 

obscura Brause, 16, 20. 

papuana Copel., 16, 21. 

pectinata, 16. 

pectinata var. brevipinnula Ros., 20. 

repens (Bory) Thwaites, 16, 20. 

rhombifoliolata v.A.v.R.. 15, 19. 

rigida J. Smith, 16, 19, 20* 21. 

roemeriana Ros., 15, 17. 

Rosenstockii Brause, 18. 

Schlechteri Brause, 15, 17. 

Schultzei Brause, 16, 20, 21. 

sepikensis Brause, 16, 22. 

sessilis Copel., 15, 16, 17. 

sinxiato-crenata v.A.v.R. 16, 20. 

stolonifera Mett., 16, 22. 

tenuifolia Blume, 15, 17. 

trichophylla Copel., 17. 

tricrenata Baker, 15, 19. 

Versteegii (Christ) v.A.v.R., 16, 18. 

Werner! Ros., 16, 21. 

WoUastonli V.A.V.R., 17. 
Livistona rotundifolia, 46. 
Lomagramma J. Smith, 401. 

angustipinna (?opel., 401, 402. 

Brooksii, 401. 

lomarioides (Blimae) J. Smith, 401. 

novoguineensis (Brause) C. Chr., 401. 

sinuata C. Chr., 401. 



Lomaria* 225. 

iatinscula V.A.V.R., 226. 

rei;t>ltita v.A.v.R., 226. 
Lomariopsis F^, 400, 

Qochinchinensis F4e, 400. 

Klngli (Copd.) Holttum, 400. 

intermedia (Copel.) Holttum, 400. 

subtrifoliata {Ck»pel.) Holttum. 401. 
l^oxoscaphe, 219, 221. 

Li3pea tranqnebarica H, MiIne«.Edwards, 99, 
102, 103. 


Ma. 168. 169. 

Mabayiff, 188. 

Madamdama, 177. 

Magydaris, 294. 

Haka, 169, 174. 

Makamanmand, 175, 188. 

Makamand, 175, 188. 

Makanika, 174. 

Makasasmamano, 175. 

Makaysa, 175. 

Maki, 168. 169, 171, 203. 

Makin, 179. 

Malpisliiaceee, 295. 

Mami. 191. 

Mamin. 191. 

Mamin-adti, 176. 

Maminpln, 191. 

Maminpisdn. 191. 

Mamosain (S. serrata), 96. 100, 101, 104, 

107, 108. 
Man, 185, 189. 198. 199. 
P. WEST. Analysis of Manila elemi, 
Manen, 175, 189. 
Maiigniffi. 202. 
Manipud, 202. 
Manungpal, 203. 
Mara, 169. 
Masans&n. 176. 
Meffarhinus. 46. 
Menippe mercenaria. 334. 
Meniscium beccarianum CesatI, 460. 

cuspidatum BItime. 444, 458. 

triphyllum SwartK. 459. 
Menispermaeec?, 294. 
Met, 196. 
Met la, 196. 

ISensr, 196. 
Microcala. 59. 65. 
Mierolepla Presl. 11, 13. 

melanorhacbis Rc»., 13. 

pilotisla, 14. 

psendohirta Ros.. 13. 14. 

scaberula, 14. 

speluncte (L.) Moore. 13, 14. 

strigosa (Thmnb.) Presl. 13. 

trichosticha J. Smith. 18. 14. 

trichosticha J. Smith var. crlabrata 
Prantl., 14. 
Momordica cochinchinensis Sprensr, 299. 
MoncMBome. 8 IS. 
Musa textilis N^. 355. 
Musacese. 855. 
Myrrhis, 294. 

Nagr. 170. 
Naffist&y. 187. 
Nasistay&n. 187. 
Nai . . . an, 170. 
Nain ... an, 170. 
Naka, 170. 
Nalab^s, 189. 
Nalibit, 198. 
N&man, 184, 185. 
Kafifirrdna, 186. 
Nap&lalo. 189. 
Nepenthicola. 60, 65. 
Nephrodium echinatum, 445. 

pteroides, 445. 

simulans Baker, 424. 

tuberculatmtt Cesati, 428. 

viscosum Baker. 426. 
Nephrolepis Schiz.olomse v.A.v.R., 24. 
»«a, 184. 186. 188, 192, 198, 
l^sad. 194. 
SgfimAn. 184. 185. 
f^S'amin. 184. 
fJganngini. 177. 

di. 187. 

sa&n. 187. 
^firanid. 193. 
»gatfi. 197. 
«Kay. 184. 
Nl, 179. 
No. 172. 183. 
No mamins&n, 188. 
Notholaena hirsuta (Poiret) Desv., 36. 


Odontosoria decipiens (Cesati) Christ (?), 

Versteegii Christ, 18. 
Oenanthe, 294. 
Oenotrichia Copel., 14. 

novsB-Guineae (Res.) Copel., 9, 14. 
Onychium Kaulfuss. 86. 39. 

siliculosum Desv. C. Chr.. 39. 
Ophichthyidae, 152. 

Orthiopteris acuminata (Ros.) Ck)pel., 8, 
9. 10. 14. 

caudata Copel.. comb, nov., 8. 9. 

cicutarioid^i (Baker) Copel., comb, nov., 
8. 9. 
Orthiopteris Copel.. 7. 

fenilacea Copel.. 7. 8. 9. 

minor (Hooker) Copel.. 8, 9, 10. 

trichophylla, Copel, 8. 
Orthopodomyia, 46. 



Pa, 169. 

Pa . . . en, 169. 

Pachyrrhizua erosus (Linn.) Urb., 145. 

Pada, 186. 

Faesia St Hilaire, 28. 

lamiana V.A.V.R., 28, 29. 

radnia (Baker) C. Chr., 28. 29. 

scabenila, 29. 
Pagr, 169. 
Paga, 170. 
Pagamoan. 188. 
PAGAYON, ADELA U. see Baisas and 

Pan, 203. 

Paiig , . . em, 169. 
Parabanhawin (S. serrata), 101. 
PARAS, ERNESTO M. Blood-chemistry 
studies in leprosy. III. Total calcium, 
diffusible and nondiftusible, calcium, 
albumin and grlobulin, 73. 
Pasardy, 177. 
Pasig, 186. 
Pati, 204. 

Patin&yon. 174, 176. 
Pay, 192. 

Pellaea falcata, 88. 
Pellaea Link, 38. 

Zippelii (Miquel) Baker, 38. 
Pertunus serratus Ruppel, 103. 
PESIGAN, T. P. Marcos A. Tubangrui: 

in memoriam, 367. 
Pi . . en 170. 
Pili, 189. 
Piman, 194. 
Pin . . . en, 170. 

Pieocnemia membranacea Bedd., 414. 
Polypodiimi cheilanthoides Baker, 437. 

inffens Brause, 419. 

invisum Forster, 457. 

pteroides Retz., 457. 
Polystichum Roth. 390. 

aculeatum (L.) Schott, 392, 398. 

alpinum Ros., 392, 396. 

Archboldii <k)pel., 892, 393.; 

bamleriantmi Ros., 392, 393. 

bolanicum Ros., 392. 894, 

Brassii Ck)pel. sp. nov., 892, 894. 

cheilanthoides Ck>pel., 392, 396. 

keysserianiun Ros., 392. 

kinabaluense C. Chr., 892, 395. 

lastreoides Ros., 412. 

lineare C. Chr.. 892, 897. 

microphyllum (Blimie) Presl, 394. 

muticum Ck>pel., 392, 394. 

Myer- Dreessii Copel., 892, 395. 

ovalifolia Ros. var. ovalifolia Ros., 394. 

papuanum C. Chr. 892, 895. 
PortunidflB Miers, 98, 102. 
Powell!. 56, 67, 68. 
Pteridacese of New Guinea, 5. 

Pteridium Gleditsch, 29. 

Aquilinum (L.) Kuhn, 29. 

revolutum (Blume) Nakai, 29. 
Pteridrys Christensen and ChinsT, 412. 

microthecia (F4e) C. Chr. et Chingr, 

. 412. 

olivacea (Ros.) Copel., 411, 412. 
Pteris Linmeus, 31. 

bambusoides Gepp, 82. 

beccariana C. Chr., 88. 

blumeana Agrardh, 81, 33. 

Brassii C. Chr., 31, 34. 

Brausei Ros., 82. 

comans Forster, 81. 85. 

cretica Linn., 31, 32. 

deltoidea Copel., 31, 32. 

ensiformis Burm., 31. 33. 

excelsa Gaud., 31, 83. 

gardneri (F^) Hooker, 31, 33, 

Keysseri Ros., 31, 34. 

lanigera Blume. 29. 

ligulata Gaud., 31, 33. 

macrophylla Copel., 81, 34. 

moluccana Blume, 31, 82. 

montis-wilhelminaa Alston. 81, 34. 

pacifica Hieron.. 31, 33. 

papuana, 32. 

pellucida Presl, SI. 32. 

quadriaurita, 22. S3. 

quinquepartita Copel., 34. 

scabripes Wall., 81, 32. 

Schlechteri Brause, 31, 34. 

semipinnata Linn., 31, 33. 

torriceUiana Christ, 83. 

tripartita Swartz, 31, 35. 

vittata Linn.. 81, 82. 

wallichiana Agardh, 31. 35. 

Warburgii Christ, 31, 35. 
Pulos, 186. 


Rachisoura, 45, 62, 64, 67. 
RAMIRO, MARIANO P. Suitability of 
Manila hemp waste for paper making 
material, 355. 
Rasd (Ilocano). 95. 
Rumohra Raddi, 397. 

adiantiformis (Forster) Ching, 397. 
aristata (Forster) Ching, 897, 898. 
Hasseltii (Blume) Ching, 397, 898. 


Sa, 198. 

Saan. 180, 181. 

a kakigm&tan. 188. 

man, 185. 
Saccoloma caudatum Cope!., 9. 

moluccanum (Bl.) Hett. 10, 
Sadi, 178. 204. 
Sadi&y, 178. 
Sadin (o) ti. 182. 



Sasrenia intermedia J. Sm., 414. 

Sagpamins&n, 177* 

Safiga. 169, 191, 203. 

Sailgka, 169, 203. 

Safigkafigrona, 187. 

SANTIAGO DOMINGO, see Gapuz, and 

Santos, Jose Kabigting, 275. 
Sapindac^e, 296. 
Schistosoma japonieum, 368. 
Schizoloma fnligineum Copel., 24. 

Schizolomsd V.A.V.R., 24. 
Scorpaenidse, 153. 
Scylla De Haan, 98 

oceanica (banhawin), 304. 

oceanic (Dana), 95, 101, 102, 103, 301, 

804, 306, 815, 316, 817, 318, 320, 321, 
322, 329, 882, 834. 

serrata De Haan. 99, 100, 103. 
serrata (Forska!), 95, 99, 101, 301, 305. 

320, 322, 382, 834. 

serrata var. mamosain Estampador, 304. 

errata var. paramamosain Estampador, 

95, 101, 104, 105, 801, 304, 317, 818, 

820, 821, 880, 884. 
tranquebarica (Fabricins) 801, 304, 

805, 810, 812, 815. 816, 317. 818, 320, 

321, 322, 829, 880, 833, 334, 885, 343. 
tranquelmrica (Fabricius) Dana. (Pa- 

rabanhawin), 95, 99, 102, 103, 104, 

tranquebarica (parabanhawin), 304. 

tranquebarica var. oceanica Dana, 101. 
Scylliorhinidse, 151. 
Scylliorhinus rudis Pietschman, 151. 
Scylliorhinus torazame (Tanaka), 151. 
Sedum maximum Sut, 294. 
Senneratia caseolaris (L.) Engl., 108. 

pagatpat Blanco, 108. 
Serranidae. 152. 

SHER, HERMAN, see YuTUC and Sher. 
Si. 168. 

Si asino ti, 182. 
Sigud, 188. 
Sinan, 169. 
Sinin, 169. 
Sipud. 202. 
Solmitmi Angatii, 2. 
Sphaerostephanos J. Smith. 460. 

polycarpa (Blume) Cope!.. 460. 
Sphenomeris Maxon. 24. 

chusan (L.) Copel., 24. 

ovatum, 24. 

guirinianum, 24. 

retusa (Cav.) Maxon. 24. 
Stellera chamaejasme L.. 294, 295. 
Stenochlaena J. Smith, 228. 

areolaris (Harr.) Copel., 228. 

intermedia Ck>pel., 400. 

iuglandifolia Presl, 228. 

Kingii Ck>pel., 400. 

palustris (Burm.) Bedd.. 228. 

Stenolepia Van Alderwexelt Van Rosenburg, 

tristis (Blume) v.A.v.R., 391. 
Stenosemia Presl, 412. 

aurita (Swartz) Presl, 412. 
Sumangka, 170. 
Sjmgramma J. Smith, 25. 

Hookeri C. Chr., 25. 

pinnata J. Smith, 25. 

Tabanus albimidus. 383. 

albitriangularis, 383. 

bilateralis 383. 

brunipes, 383. 

ceylonicus, 888. 

factiosus 879. 

flaviventris 379. 

fiexiles, 379. 

fumifer, 379, 383. 

graiseoscutellatus, 379. 

griseipalpis, 383. 

hilaris. 883. 

immanis, 383. 

ixion. 379. 

latifacies, 383. 

malanopygatus 379. 

malayensis, 379, 383. 

minimus, 883. 

nemocallosus, 383. 

nemoralis, 388. 

partitus, 383. 

philippinensis, 379. 

reducens, 379, 380, 382. 383, 384. 

rubidus. 878, 382. 

rufiventris 382. 

striatus, 879, 380, 381, 382, 383, 284. 

univentris, 379. 

vagus, 888. 

vanecki, 383. 
Taenitis Willdenow, 25. 

blechnoides (Willd.) Swartz, 25. 

Brausei Ros., 32. 

requiniana (Gaud.) Copel.. 25. 
Taga, 202. 
Tagaand. 183. 
Tagi, 169, 170, 203. 
Tapeinnidium (Presl) Christensen, 23. 

Denhami, 23. 

longipinnulum ((Jesati) C. C3hr., 23. 

marginale Copel., 23. 

moluccanimi (Blume) C. Chr., 28. 

obttisatum v.A.v.R., 23. 

pinnatum (Cav.) C. Chr., 23. 

stenocarpimi v.A.v.R., 23. 

tenue (Brack.) Copel., 23. 
Tfipno, 190. 
Tari, 170. 
Tectaria Cavanilles, 413. 

andaiensis (Baker) C. Chr,. 413. 414. 

angulata (Willd.) Copel., 414, 418. 

bamleriana C. Chr., 416. 

Barclay! (Carr.) C. Chr.. 413, 416. 



baccariana (Cesati) C. Chr., 414, 419. 

Bryanti Copel., 419. 

eesatiana (C. Chr.) Copel., 413, 416. 

coadunata, 416. 

craspedosora Copel., 413, 419. 

crenata Cav., 413, 417. 

cristovalensis (C. Chr.) Alston, 414, 418. 

decurrens (Presl) Copel., 414, 419. 

devexa (Kunze) Copel. var. novosnii- 
neensis Ros., 418, 414. 

diversisora Copel., 419. 

ferrusflnea (Mett.) Copel., 413. 415, 416. 

ETisantea (Blume) Copel., 414. 

Rymnocarpa Copel., 413, 415. 

irregnilaris (Presl) Copel, 413, 415. 

Kingii Copel.. 413, 414. 

ledermanni (Brause) C. Chr., 418, 415. 

leuzeana (Gaud.) Opel., 413, 415. 

malayensis, 416. 

melanocaulis (Blume) Copel., 413, 418. 

Menyanthidis (Presl) Copel., 414, 418. 

minuta (Jopel., 414, 419,. 

novofifuineensis (Ros.) C. Chr., 414, 418. 

papuana Copel., 416, 417. 

pleiosora (v.A.v.R.,) C. Chr., 413, 417. 

pleocnemioidea (v.A.v.R.) C. Chr., 414. 

pubescens Copel.. 413, 415, 416. 

semibipinnata (Wall.) Copel.. 414, 418. 

siifolia (Willd.) Copel., 414, 419. 

subaequalis CTopel., 412. 

teratocarpa (v.A.v.R.) C. Chr., 413, 417. 

ternatensis V.A.V.R., 413, 417. 

vasta (Blume) Copel,, 419. 

Weinlandii (Christ) CJopel., 413, 418. 
Teratophyllum Mettenius 400. 

aculeatum (Blume) Mett., 400. 

Williamsii, 402. 
Thamnopteris, 210. 
Therapia magna sterlizans, 135. 
Thwaites C. P. 1287, 423. 
Thylacospermxun rupifragum Schrenk., 294. 
Thymelaeaceie, 295. 
ThyTsanosoria Crepp, 400. 

dimorphophylla Gepp, 400. 

pteridiformis (CJesati) C. Chr., 400. 
Tinn&ar, 174. 

Tinospora oordlfolia Miers., 294. 
Ton kaand ti, 181. 
Tonan6 ti, 181. 
T6no. 172, 
Topomyia, 46. 
Tripteroides, 44-60, 69. 
pupae and larvae, 43. 

Trypanosoma equinum. 159. 
equiperdum. 141. 
evansi, 186. 186. 137, 141, 142, 156, 157, 

164, 383. 
gambiense. 142. 156. 
rhodesiense. 141. 142. 


UICHANCO, LEOPOLDO B. A revision of 
the genus Ectenus Dallas, vdth de- 
scription of a new species (Hemip- 
tera, Pentatomidae), 285. 
Urn. 169. 

Umbellifer». 291, 294, 295. 
Unay. 187, 190. 
Uranotaenia, 46. 
Uray. 193. 

and. 181. 

inton, 181. 

(no) ipapaano, 188. 

kaan6, 181. 

no, 181. 

(no) kasano, 188. 


bigting Santos. 275. 

prepositions in Iloko, 167. 


Wen, 192. 

WEST, AUGUSTUS P., »ee Manalo and 

Woodwardia J. E. Smith, 227. 228. 

auriculata, 228. 

orientalis. 228. 

radicans (L.) Smith, 227, 228. 

unigemmata, 228. 

Yano, 183. 

YUTIXJ, L. M. Observations on the pre- 
valence of tabanid flies and surra- 
transmission experiments, 379. 

Observations on the distribution of 
Trypanosoma evansi in the bodies of 
rats and guinea pigs during different 
stages of infection, 155. 

Zeugnomyia, 46. 

Vol, 78, No. 4 

December, 1949 

The Philippine 
Journal op Science 






Published by the Institute of Science 
(Formerly Bureau of Science) 

' A. S. Arguelles, D.Sc, Editor 

Eduardo Quisumbing, Ph.D., Associate Editor 



Makuel L. Rodus, Ph.D.; F. T. Adriano, Ph.D. 

Joaquin Maranon, D.Sc; R. H. Aguilar, Ch.E. 

Patrocinio Valenzuela, Ph.D.; Marcos M. Aucante, Ph.D. 

A. J. Hermano, D.Sc; Felix V. Espino, Ch.E. 

V. Elicano, B.S.; Antonio D. Alvir, Ph.D.; Jose Feliciano, Ph.D. 

Eocperimental Medicine 

Daniel de la Paz, M.D.; Arturo Garcia, M.D.; Onopre Garcia, M.D. 

Cristobal Manai4Ang, M.D.; Isabelo Concepcion, M.D. 

H. W. Wade, M.D,; Walfrido de Leon, M.D. 

Clinical Medicine 

Antonio Sison, M.D.; Liborio Gomez^ M.D., Ph.D.; H. Lara, M.D. 
Jose Rodriguez:, M.D.; Carmelo Reyes, M.I). 


Elmer D. Merrill, D.Sc; E. B. Copeland, Ph.D.; A. F. Fischer, C.E., M.F. 

T. G. Fajardo, Ph.D.; Rafael B. Espino, Ph.D. 

Nicanor G. Teodoro, Ph.D.; Feliciano M. Clara, Ph.D. 

Nbmesio B. Mendiola, Ph.D. 


Leopoldo B. Uichanco, D.Sc; Deogracias V. Villadolid, Ph.D. 

Heraclio R. Montalban, M.A.; Gonzalo Merino, Ph.D. 

Canuto G. Manuel, D.Sc; Manuel D. Sumulong, M.S., D.V.M. 

l^PB M. YuTUC, D.V.M. ; Faustino Q. Otanes, M.S. 

Leopoldo S. Clemente, Ph.D. 

H. O. Beyer, M. A. ; Ricardo E. Galang, M.A. 

(Bi^ered mt tb€ Post Office at BSanila, Philippines, as s^ond-class matter) 


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